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The philosophical basis of biological classification.

Review of Marc Ereshefsky (ed.) The Units of Evolution

Article  in  Studies In History and Philosophy of Science Part A · April 1994

DOI: 10.1016/0039-3681(94)90032-9

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 ESSAYREVIEW

The Philosophical Basis of Biological

Classification
John Da-put*

Marc Ereshefsky (ed.), The Units of Evolution: Essays on the Nature qf Species
(London: MIT Press, 1992) xvii + 405 pp. ISBN o-262-05044-7 Cloth E44.95,
ISBN o-262-55020-2 Paperback E24.75.

THE NATURE of biological species raises a variety of important philosophical

questions in both the philosophy of science and metaphysics, not to mention
central areas of biology. This volume brings together an excellent collection of
the key papers bearing on these issues written in the last thirty years both by
biologists and by philosophers. This will increase the accessibility of these
debates to scholars from a variety of disciplines, and for specialists and
teachers in the area it will be a great convenience. For these reasons it is to be
warmly welcomed.
One source of interest in questions about species is the very fact that it is an
area in which scientific and philosophical interests converge. A disappointing
aspect of the anthology is that Ereshefsky adopts an essentially apartheid
division of the essays: the first part of the book contains essays by biologists,
the second (with the exception of two papers by Ghiselin) essays by writers
known primarily as philosophers. One cannot help suspecting that this reflects
the drearily deferential attitude of philosophy towards science so characteristic
of contemporary philosophy of science. Ereshefsky writes in his general
introduction: ‘Philosophers often strain to be practical; biologists enjoy a
relevant philosophical discussion’ (p. xvi). Perhaps, one is tempted to add,
after work and over a pint of beer. 1 carp on this point because it seems to me
that one of the most striking impressions that emerges from the papers in this
collection is the deep interconnectedness of biological and philosophical issues.
It is a pity that this anthology does not do more to highlight these connections.

*Department of Philosophy, Stanford University, CA 94305-2155, U.S.A.

Stud. Hist. Phil. Sri., Vol. 25, No. 2, pp. 271-279. 1994.
Elsevier Science Ltd
Printed in Great Britain
Pergamon 0039-368 l/94 $7.00 + 0.00

271
212 Studies in History and Philosophy of Science

The most fundamental issue about species can be found at the very begin-
ning of the book.’ The juxtaposition of the title ‘The Units of Evolution’ with
the subtitle ‘Essays on the Nature of Species’ might naturally suggest to the
casual reader that species are, indeed, the units of evolution-an assimilation
made explicit on the first page of the general introduction (p. xiii).2 But this is a
highly contentious issue. Two questions arise immediately: first, whether or
why species really are coextensive with units of evolution; and second, what,
anyhow, is a unit of evolution?
Beginning with the second of these questions, it is a familiar enough
observation that species are what evolve. Given only the assumption that
evolution involves the gradual replacement of organisms by organisms with
slightly different properties, as opposed to changes in particular organisms, it
can be said that what evolves is a group rather than an individual organism.
But clearly more is at stake than simply a change in the distribution of
properties of organisms within some group of organisms: if it were just this,
then every higher taxon, being composed of evolving species, would itself be a
unit of evolution. A more focused conception of the units of evolution requires
an account of a way in which certain groups of organisms evolve in some
coherent way. More specifically, a view suggested by a number of authors in
this anthology is that species provide the boundaries within which evolutionary
changes are constrained. The classic such account is the so-called biological
species concept, which defines a species as a group of organisms connected by
reproductive links and reproductively isolated from members of all other
species. Genetic novelties can eventually reach all the descendants of the
members of the species by means of reproductive links, but have no path to
any other organism.
The first part of the book is organized in terms of various critiques of the
biological species concept just introduced. Though this concept is appro-
priately introduced by Ernst Mayr, the biologist most closely associated with
its defence, it is perhaps a little unfair that Mayr’s contribution takes the form
of a rather brief extract from a classic, but 30-year-old, expository text. This
comes across as very dogmatic when set against the more recent and self-
consciously contentious papers that follow. (In fact, the most robust defence of
the biological concept can be found in the penultimate paper in the volume, by

‘It has become a convention in this field to make much of a distinction between the species
category and species taxa. the various particular species such as Homo sapiens. Rulrus ra~tus, etc.
Thus Ereshefsky: ‘The term species refers to two types of entities:. Species /UYUare groups of
organisms, Dog is a species taxon and chickadee another. The species rcuegnry, on the other hand.
is the class of all species taxa’ (p. xiv). Similarly, I suppose, the term dog is equally ambiguous. Dog
individuuls are particular organisms (such as Fido). and the dog ccctegor_v is the class of all dog
organisms. It has never been clear to me what dreadful confusion was to be avoided by the almost
religious insistence on this dichotomy; whatever it is. 1 shall expose myself to the risk in what
follows.
‘Though the opposite is a thesis of Ereshefsky’s own contrrbution to the anthology.
Biological CIassiJcation 273

Ghiselin.) A more general difficulty with this part of the book is that somewhat
at cross purposes with this focus on the biological species concept is the
problem of providing an illuminating taxonomy of theoretical accounts of
species. Ereshefsky notes (p. 3) that seven alternative definitions of a species
occur in the readings. Later he distinguishes (p. 9) between three main schools
of systematics, but without attempting to relate these to the various defini-
tions. And finally, Mayr, in the excerpt just mentioned, locates the biological
species concept in a quite different tripartite division, distinguishing it from
typological (essentialist) and nominalist concepts (pp. 16-17). I suspect that at
this point the naive reader might be quite confused.
We may begin by noting that none of the critics of the biological species
concept is naturally described as a strict essentialist or strict nominalist,
though contemporary morphological taxonomists, stigmatized by Mayr as
essentialists, do perhaps flirt with nominalism. More relevant here is the
tripartite distinction mentioned by Ereshefsky, who distinguishes cladism,
phenetic taxonomy, and evolutionary taxonomy. The basic ideas behind the
first two are relatively straightforward. Phenetic, or morphological, taxo-
nomists are concerned to identify degrees of overall similarity between organ-
isms. In its recent guise of numerical taxonomy it appeals to sophisticated
techniques for analyzing large numbers of traits. This position is represented in
the anthology by the paper by Sokal and Crovello. Cladism, represented in
rather different versions by Wiley, Cracraft, and Mishler and Donoghue,
insists on the phylogenetic coherence of taxonomic units, requiring that such
units contain all and only the descendants of some ancestral population.
(Technically speaking, taxonomic units must be monophyletic.) Evolutionary
taxonomy is harder to pin down.3
Like the biological species concept, its best known representative, evolu-
tionary taxonomy has succeeded in a somewhat imperialistic name grab.
Certainly it is no more peculiarly concerned with evolution than is cladism.
One point of difference is that evolutionary taxonomists do not generally insist
on monophyly. Thus, for example, they can deny that birds are a kind of
reptile. Of more present significance is that whereas cladism is exclusively
concerned with the overall pattern of evolution, as displayed in a phylogenetic
(or genealogical) tree, evolutionary taxonomy also addresses the process of
evolution. With these distinctions in mind, we can sort out some rather
different criticisms of the biological species concept. In addition, we can see
these criticisms as implicitly reflecting different conceptions of the appropriate
relation between theory (in this case evolutionary theory) and classification.

‘As I interpret these various concepts, Wiley’s ‘evolutionary species concept’ is not a version of
evolutionary taxonomy.
274 Studies in History and Philosophy of Science

To oversimplify somewhat, we may say that phenetic taxonomy aims to

provide a maximally theory-free classification. No doubt taken to an extreme
this is incoherent. There is no such thing as an absolute degree of similarity, if
only because any two objects will always have infinitely many shared proper-
ties and infinitely many distinct properties. Some conception of what traits
matter is inevitably involved in applying any kind of phenetic taxonomy.
Nevertheless, with this acknowledged, the goal of taking as much of what
might matter as possible into account in discerning similarity seems an
intelligible one, and intelligible specifically as a way of remaining as theory-
neutral as possible. Sokal and Crovello also argue that attempting to be less
theory-neutral, at least in the manner of the biological species concept, is
incompatible with sound empiricism. Reproductive connections and barriers
are, they argue, almost inevitably inferred, in practice, from estimates of
morphological similarity.
Cladistics, on the other hand, is committed to the attempt to produce a
classification that reflects evolutionary history, and traits are important just to
the extent that they reveal phylogeny. The ideal cladistic taxonomy provides
names of taxa in one to one correlation with forks in the phylogenetic tree. To
pick up a question left hanging some pages back, cladistics can be seen as the
school of systematics that takes the thesis that species are units of evolution
(though not necessarily vice versa) to be a matter of definition. Cracraft
(p. 103) cites this at the beginning of a list of advantages for his phylogenetic
species concept, and points out that it settles conceptually various vexed
questions over, for example, the relations between species and subspecies.
The various positions that can be grouped together under the heading of
evolutionary taxonomy agree that evolutionary change flows down phylo-
genetic channels, but consider that an adequate definition of species must
specify what makes a group of organisms such a conduit for evolutionary
change. Thus by contrast with the case of phenetics, in which species are
merely conceived as classificatory units, and coincidence with evolutionary
units would be purely fortuitous, and cladistics, in which this coincidence is a
matter of definition, for evolutionary taxonomy the coincidence is an empirical
hypothesis: the hypothesis that the feature which is used to define species is in
fact the feature that makes a group of organisms a possible conduit of
evolutionary change. Given this empirical commitment, it is no surprise that
the various evolutionary species concepts should be more of a hostage to
empirical fortune than the other two just mentioned. And thus advocates of
rival evolutionary species concepts deploy various empirical claims about the
distribution of organisms and its causes to argue for the inadequacy of the
biological species concept.
The biological species concept develops the claim that what separates
channels of phylogenetic descent is reproductive isolation. As developed by
Biological Class$cation 275

Mayr this also encompasses a theory of the origins of particular species in the
geographical isolation of peripheral populations, which are thereby enabled to
evolve independently, thus eventually generating divergence sufficient to make
interbreeding with the original population impossible. In a perfect world, we
might hope that every phylogenetic division involved the establishment of
reproductive isolation, and all separate lineages were reproductively isolated
from each other. If, moreover, there were sharp morphological distinctions
between all lineages, we could suppose that cladism, pheneticism, and the
biological species concept would all coincide on the same divisions. Something
like this is suggested by Michael Ruse in his essay in Part 2. However, the clear
consensus of the authors in Part 1 is that things are messier than this, and
messy in ways that point to serious limitations to the applicability of the
biological species concepts.
The friendliest amendment to the biological species concept is that of
Paterson, who suggests that the standard concept involves a naive conception
of the process by which gene flow is restricted. His ‘recognition’ concept
proposes that speciation is achieved when a group of organisms develops a
way of identifying conspecifics as mating partners. Though some important
differences from the isolation concept emerge from this proposal, it is surely in
the same spirit. More radical objections begin with the classic difficulty of
accommodating asexual species. A great many organisms do not engage in any
form of sexual reproduction, and are thus isolated from all other organisms.
Mayr (p. 23) notes that this precludes the application of the biological species
concept and remarks that species in these cases are customarily delimited on
the basis of morphological difference. Ghiselin (p. 373) asserts more bluntly
that such groupings must be thought of as no more than ‘pseudospecies’.
Whereas asexual organisms are reproductively connected to too few organ-
isms, other organisms are connected to too many. Indeed hybridization
between apparently distinct species is extremely common. One classic case is
that of North American oaks as discussed in the paper by Van Valen. There is
apparently a great deal of gene flow between several species of oaks, but it also
appears that morphologically sharply distinct species have coexisted for
millions of years. Again Ghiselin (p. 371) is happy to bite the bullet and
suggest that such groups of oaks may simply turn out to form one (polymor-
phic) species. However, Van Valen, as also Ehrlich and Raven, draw a
different conclusion. The fact that both in asexual species (as Mayr, p. 23
concedes) and in groups of species with considerable interspecific gene flow
there are often morphologically well-defined species provides a strong argu-
ment for thinking that something other than reproductive isolation is capable
of maintaining taxonomic discontinuities and hence, very probably, cohesive
channels for evolutionary change. The force that these authors propose is
ecological: specific differences are maintained by selective pressures.
216 Studies in History and Philosophy qf Science

Ehrlich and Raven reach their conclusion not so much from considering
interspecific gene flow as by emphasizing the limitations of intraspecific gene
flow. They cite a range of evidence that within many species gene flow is
extremely limited and local. Again this observation suggests that some source
other than reproductive isolation maintains taxonomic distinctness, and the
obvious candidate is a common selective regime. In sympathy with the idea
that it is groups delimited by gene exchange that constitute the units of
evolution, Ehrlich and Raven conclude that the units of evolution are generally
not species but local interbreeding populations.
These various cases do not complete the picture of taxonomic diversity. At
one extreme there are highly diverse genera in which there is a great deal of
gene flow and few, if any, clear divisions. Van Valen (p. 73) suggests that these
might be cases of taxa in which there were no species, though considering his
example of the genus Rubus, that is certainly a case where we would want to
make distinctions (e.g. between blackberries and raspberries) on more practical
grounds. At the other there is a range of cases that shade into asexuality, of
organisms that are very largely self-mating (Templeton, p. 164). (This case falls
between full asexuality and the localized gene exchange emphasized by Ehrlich
and Raven.)
A natural attempt to impose some order on this situation is Templeton’s
cohesion species concept. The cohesion of a group of organisms, that is, their
continuity as morphologically distinct from other organisms, can apparently
be brought about in various ways. Templeton mentions gene flow, natural
selection, and ecological, developmental and historical constraints as among
the forces that can promote such cohesion. Although this concept surely
resolves many of the problems raised by criticisms of the biological species
concept, the price may seem high. No general account of speciation survives in
Templeton’s concept. Indeed one may reasonably see this as coming very close
to a phenetic concept: species are whatever groups can be clearly distinguished
from related or similar groups, and we should pay attention to the various
mechanisms that promote such distinctions in determining the attention we
accord to particular phenetic features. And it is surely a strong empirical claim
that all these sources of division equally define units of evolution. Given Sokal
and Crovello’s undoubtedly correct emphasis on the necessity of phenetic
analysis in the application of any species concept, and the empirical diversity in
the patterns of variation that emerge from the biological data, perhaps this is
just where we should expect to end up. Or rather, perhaps what we are left with
is a clear choice between a basically phenetic approach to taxonomy, though
stripped of the theoretical innocence generally associated with the numerical
version, or an explicitly phylogenetic, probably cladistic, concept. And this
rather simpler division brings us back to the question: Why should our units of
classification coincide with the units of evolution?
Biological ClassiJication 277

I shall approach this question by way of a brief survey of the more

philosophical part of this volume. The first three of these papers provide
important philosophical background to the problem of species, and might well
be read ahead of the more technical biology in Part 1, certainly by a non-expert
reader. The papers by Hull and Sober analyse essentialism and the sense in
which it has been refuted in biology by the general acceptance of Darwinian
evolution. Though the paper by Hull is widely cited, on rereading it seemed to
me that the decision to reprint the second half (the paper was originally
published in two parts) was questionable. The first half, on essentialism, is still
valuable, but the second, in which he attempts to provide a disjunctive
definition of species, seems a bit dated. Hull has done a great deal more work
on this topic since 1965, and I doubt whether he would want to defend many of
the details of this account now. For example, the idea of determining a unit of
taxonomic length as ‘the morphological distance usually indicative of inter-
breeding status among contemporary organisms’ is difficult to reconcile with
the more recent empirical results in Part 1. The third paper in this section, by
Beatty, concerns the way Darwin himself used the term ‘species’, specifically
his strategy for dealing with the ways in which the species concept he was
forced to use embodied much of the theory he was concerned to refute. This
paper has important implications for understanding theory-change and the
theory-ladenness of scientific concepts, and it provides a historical perspective
on what I have suggested remains an important issue in understanding species.
The next two papers introduce one of the most contentious ideas in recent
philosophy of biology, the idea that species, far from being natural kinds, are
not kinds at all, but individuals. This thesis was introduced by Ghiselin and
has been defended in detail by Hull, and appropriate papers by these authors
are presented. Although the arguments are quite complex, the only point I
should stress here is that central to this issue is the question whether species are
to be seen as primarily units of classification or units of evolution. It is the
insistence that they are the latter, suggesting that they are not kinds that figure
in biological laws, but the objects that fall under such kinds, that primarily
motivates the thesis that species are individuals.
Opposing views are offered by Kitcher and Ruse. Ruse defends the tradi-
tional idea that species are indeed natural kinds, but not in the sense of
possessing unique essential properties, but in the sense (often associated with
the name of William Whewell) of being sets on which various differently
motivated principles of classification coincide. However, as noted above, this
claim of convergence seems excessively optimistic in view of the empirical
evidence. Kitcher argues that species are neither individuals nor natural kinds,
but merely sets. Moreover, because he does not believe that different principles
of classification necessarily coincide on the same sets, he defends a pluralistic
conception of species. That is to say, he rejects the assumption that there must
278 Studies in History and Philosophy of Science

be some unique correct answer to the question of what is the appropriate

criterion of species membership. Most generally, he insists that both structural
and phylogenetic principles of classification may be relevant for various
purposes. Though there are some very attractive consequences of such a
position, it is liable to encounter strong resistance on the basis of intuitions
about the fundamental unity of science, and the related accusation that such
pluralism threatens to undermine the possibility of meaningful communication
between scientists.4
The kind of pluralism advocated by Kitcher must be sharply distinguished
from that advocated by Mishler and Donoghue and by Ereshefsky. These
authors urge that although we should insist on phylogenetic coherence as a
condition of adequacy for biological taxa, how much of the genealogical nexus
we want to distinguish, and on what grounds, may vary greatly from one area
of biology to the next. Ereshefsky draws a number of conclusions from this
position. One, which is surely correct, is that the distinction between species
and higher taxa, often taken to be fundamental, becomes quite murky.
Ereshefsky argues, in fact, that not only species, but also many higher taxa,
may be units of evolution. This idea brings us once more to the contrast
between units of evolution and units of classification. One view I have
mentioned is that species are, by definition, the units of evolution. This has the
drawback that some species might prove to be very large (huge polymorphic
species of oaks, or the whole genus Rubus, or even, on Ereshefsky’s view, taxa
currently located at any higher level) and some may be tiny. From the point of
view of classification one would like the basal units to be rather more
homogeneous, especially if, as these pluralistic cladists believe, there is no
principled grounds deriving from phylogeny for privileging as species any
particular subset of the monophyletic segments of the genealogical tree. But
once this divorce between units of evolution and of classification is acknow-
ledged, it becomes difficult to see why we should stop short of the more radical
pluralism advocated by Kitcher. After all, a classification should surely be of
use for areas of biology other than the study of evolution.
I shall conclude with a few final thoughts on the relationship between science
and philosophy as it appears in this issue. Obviously enough, there are
empirical facts that are fundamental to these issues and with respect to which
we should surely defer to empirical science. Essentialism, I suppose, might
have turned out to be true; species might have been perfectly isolated from one
another, or there might have been unrestricted gene flow within all species;
there might not have been any asexual organisms; and so on. On the other

4The thesis of the unity of science is criticized at length in my book The Disorder of Things:
Metaphysical Foundations of the Disunity of Science (Cambridge, Mass.: Harvard University Press,
1993). There I also defend a pluralistic view of species similar to that of Kitcher.
 Biological CIassiJication 279

hand, as I hope has been clear from the preceding discussion, questions about
species are permeated by paradigmatically philosophical problems. Although it
is surely an area of science in which practitioners and professional philoso-
phers have had more interaction than most others-if only because of Hull’s
role as a mediator between the two disciplinary sides-it seems to me that
more is needed. I am struck by the very mixed level of philosophical sophistica-
tion in the discussion of the species question by biologists. Sometimes it seems
to be supposed that the question of which species concept is correct is simply a
matter for empirical discovery; and there is, perhaps consequently, little
discussion of the question: What is the ultimate purpose of identifying organ-
isms as members of species? (Indeed, this is a question only touched on in
passing in the present anthology.) Philosophers need to recognise that they
have just as crucial a professional expertise in this area as do biologists. No
doubt biologists could also accuse philosophers of biological naivete, though
that is for them not me to decide. At any rate, perhaps what I referred to as the
apartheid division of this volume is a reasonably accurate symptom of the way
things currently stand. A more imaginative presentation of the material in this
collection might, however, have done more to show the importance of bringing
the two sides together.

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