Species Concepts and the Ontology of Evolution

JOEL CRACRAFT

Departmentof Anatomy
University of Illinois
Chicago, Illinois 60680,
U.S.A.

and

Field Museum of NaturalHistory

Chicago IL 60605
U.S.A.

ABSTRACT: Biologists and philosophers have long recognized the importance of species,
yet species concepts serve two masters, evolutionary theory on the one hand and taxonomy
on the other. Much of present-day evolutionary and systematic biology has confounded
these two roles primarily through use of the biological species concept. Theories require
entities that are real, discrete, irreducible, and comparable. Within the neo-Darwinian
synthesis, however, biological species have been treated as real or subjectively delimited
entities, discrete or nondiscrete, and they are often capable of being decomposed into
other, smaller units. Because of this, biological species are generally not comparable across
different groups of organisms, which implies that the ontological structure of evolutionary
theory requires modification. Some biologists, including proponents of the biological
species concept, have argued that no species concept is universally applicable across all
organisms. Such a view means, however, that the history of life cannot be embraced by a
common theory of ancestry and descent if that theory uses species as its entities.
These ontological and biological difficulties can be alleviated if species are defined in
terms of evolutionary units. The latter are irreducible clusters of reproductively cohesive
organisms that are diagnosably distinct from other such clusters. Unlike biological species,
which can include two or more evolutionary units, these phylogenetic species are discrete
entities in space and time and capable of being compared from one group to the next.

KEY WORDS: Species, evolutionary theory, individuality

I. INTRODUCTION

There exists within contemporary evolutionary biology a mild compla-

cency toward concepts of species and hypotheses about their origin. For
most biologists whose research programs are designed to elucidate evolu-
tionary process-level phenomena, the biological species concept is firmly
entrenched and speciation is assumed to occur primarily via geographic
isolation of small peripheral, founder populations. Historically, these views
appear to have gained acceptance because of the influence of Theodosius
Dobzhansky (1937) and Ernst Mayr (1942, 1963). The widespread con-

Biology and Philosophy2 (1987) 329-346.

© 1987 by D. Reidel PublishingCompany.
330 JOEL CRACRAFT

tentment that evolutionary biologists have with the biological species

concept (hereafter, the BSC) is surprising given the voluminous and
contentious literature that has persisted for decades. Despite the BSC
having come under strong criticisms from many quarters, most of the
arguments have had little impact within evolutionary biology. Philoso-
phers, likewise, have generally adopted the BSC and have paid scant
attention to the controversies within the systematic literature (Rosenberg
1985 is a notable exception). Indeed, until the introduction of the "species
as individuals" proposition (Ghiselin 1974, 1981; Hull 1976, 1977, 1978,
1980, 1981a), philosophers of biology have contributed relatively little to
the debate over species concepts. Importantly, also, discussion about
"species as individuals" has been almost entirely within the context of the
BSC. Because of this, some critical philosophical and biological problems
within evolutionary theory have gone unexplored.
This paper addresses the inseparable relationship between species
concepts and evolutionary theory. It attempts to shift emphasis away from
species individuality (see papers of Mayr, 1987, and Ghiselin 1987, and
ensuing discussions) - but without diminishing the importance of that
concept - and toward certain biological and philosophical issues which
have received too little attention.

II. OPERATIONALISM AND SPECIES CONCEPTS

Can species be defined by theory-independent criteria or must a definition

rely upon a prior committment to a particular theory of the evolutionary
process? To some evolutionary systematists, such as Bock (1977, p. 873),
the definition of species "stands for the concept of the species in evolu-
tionary theory" (in this case, classical neo-Darwinism) and need not -
indeed probably should not (e.g., 1977, p. 874) - include mention of
criteria that might be used to recognize entities that could be called
species. Rosenberg (1985, pp. 182-187) also rejects a strict "opera-
tionalist" definition of species, but recognizes the need for defining species
independent of any particular theory. Species, after all, are the "phe-
nomena that evolutionary theory can be expected to explain and that can
be expected to test the theory" (Rosenberg 1985, p. 182). To the extent
that this is true, it might be argued that the chosen species concept must
be largely independent of any particular version of evolutionary theory
(and here speciation can also be included). It is not apparent, however,
that this approach to species concepts will ever be fruitful. The organisms
of the world could be partitioned into clusters (call them species) in many
different ways depending upon how one chose to do the partitioning.
Without external criteria referenced to some theoretical or belief system,
rationale choice among the competing species concepts would be impos-
 SPECIES CONCEPTS AND THE ONTOLOGY OF EVOLUTION 331

sible. Definitions always include undefined terms, many of which have

reference to theoretical statements, and thus no definition can ever be
entirely theory-neutral. But this is not the same thing as saying we should
prefer "nonoperational" definitions of species. To the scientist, definitions
must ultimately be useful empirically no matter how the structure of that
definition might be characterized. One definition is preferable to another
because it allows us to understand nature better, that is, within the context
of theories and hypotheses about biological patterns and processes, that
definition lends more coherence and explanatory extensibility to those
conjectures. Such will be the thrust of some of the following discussion
about species concepts.

III. INDIVIDUALITY AND DISCRETENESS

Philosophers and biologists may have various reasons for arguing that
species are (or are not) individuals, but unless these arguments have
empirical consequences regarding the manner in which biologists view
species or processes involving them, the arguments themselves will have
little, if any, significance for biology. In fact, at least two components of
the debate over species individuality seem to have measureable empirical
consequences for evolutionary biology.
First, the individuality debate directs attention to questions about the
logical structure of theories about speciation and evolutionary processes,
particularly with regard to the identification of the entities included within
the domains of these theories (Ghiselin 1987). If theories are conjectures
describing the behavior of collections (classes) of individual entities, it
is obviously of some importance to know what individuals are being
addressed by the theory in question. Moreover, the mere description of
nonrandom pattern requires the correct individuation of entities. If species
are not viewed as individuals, then what role can they play in evolutionary
theory? If species are not the entities of evolutionary theory, what are?
And if species are not individuals, can we even speak about "the origin of
species"? The problem of "species as individuals" is only one example of a
pervasive problem within evolutionary biology, namely a general lack of
concern over questions about ontology. Many "entities" within biological
theory - "niche," "adaptive zone," "community," a "paraphyletic/poly-
phyletic" taxon, to name a few - have questionable ontological status, and
as such imply a lack of precision in the formulation of the theories
themselves.
The second saluatory manifestation of the debate over individuality is
to raise the issue of the discreteness of species and the means we have of
delineating them through space and time. Such issues have direct empirical
consequences for biology, not only with respect to biologists' efforts to
332 JOEL CRACRAFT

catalogue nature's diversity but also to the validity of our current concep-
tions about the evolutionary process. Indeed, conflicting opinions about
the discreteness of species is a major problem for the conceptual unifica-
tion of the so-called neo-Darwinian synthesis, a problem that is perhaps
partially resolved by considerations of species individuality. The issue of
individuality, moreover, is at the heart of arguments of those evolutionary
biologists who see a need to view the patterns and processes of evolution
as being hierarchically arranged into microevolutionary and macroevolu-
tionary levels (Eldredge and Cracraft 1980; Vrba and Eldredge 1984;
Vrba and Gould 1986).

IV. SPECIES CONCEPTS AND THE REQUIREMENT OF THEORY

Biologists, and philosophers (e.g., Rosenberg 1985, Chapter 7), think

species are important because they provide an ontology for a description
of nature's diversity and for processes postulated to operate in nature (e.g.,
speciation, competition, and so on). Scientific conjectures (laws, theories,
hypotheses), at least if they are to have any measure of generality, do not
pertain to individual entities, which are spatiotemporally bounded, but to
classes of these entities that are, at least in principle, spatiotemporally
unrestricted (Hull 1974, pp. 47-48; Ghiselin 1987). This distinction
between classes and individuals and their different roles in the structure of
theories has clearly been the impetus behind the writings of Hull and
Ghiselin on species individuality. Some adverse reactions within the
biological and philosophical communities to species individuality seem to
have missed this point. Many biologists, perhaps, can see little or no need
in their daily scientific practice to worry about whether species are or are
not individuals, and thus dismissing the issue as unimportant seems easy
enough'. But Ghiselin and Hull, I think, are correct: species individuality
is of fundamental importance. The confused ontology about species within
the neo-Darwinian synthesis is ample demonstration of this (sections V
and VII).
The "neo-Darwinian synthesis" is less a coherent theory of evolution
than a loosely tied package of conceptualizations or hypotheses describing
change within systems ranging from molecules to higher taxa. As a con-
sequence, these theoretical statements include a wide variety of ontologies.
For many biologists "evolution" is change in gene frequency, in which case
the units or entities under consideration are "genes." Other biologists see
evolution in terms of the origin of taxa, thus "species" become the
currency of the theoretical statements and "genes" might be ignored
altogether. The "units of evolution" that one recognizes therefore depend
upon how one conceives of "evolution" and the "hierarchical level" to
which theoretical statements pertain 2.
 SPECIES CONCEPTS AND THE ONTOLOGY OF EVOLUTION 333

This paper is concerned with species and processes such as speciation

that employ species as their units of change. Despite this restricted focus,
we can ask whether the logical structure of theories in general places
constraints on the nature of the entities contained within those theories. If
so, then an understanding of those constraints should tell us something
about theories, such as speciation, within which ontological issues are such
a matter of contention. Unfortunately, the relationship between theory
structure and the entities of those theories seems to have been insuf-
ficiently explored. One such discussion, however, is that of Gaukroger
(1978) who recognizes (p. 15) explanatory structures as consisting of "an
ontology [i.e., the entities of the explanatory discourse], a domain of
evidence, a system of concepts relating these two, and a proof structure
which specifies the valid relations which can hold between the concepts of
the system." We are concerned here with the characteristics that might be
expected of these entities. Based on Gaukroger's (1978, pp. 39-45) more
general treatment, I suggest four are important for any discussion of
species and the theories that use them.
1. Reality. We require entities to have some "reality" or existence
though they might be unobservable, even in principle (as in some physical
theory). The notion of reality is defined, or required, by theory. The
problem of observability/nonobservability is present even in evolutionary
biology, although perhaps not to the extent as in the physical sciences.
Thus, despite the claims of some biologists, species themselves are in some
sense nonobservable but that does not mean we should abandon the
notion of species reality (see also Rieppel 1986). If species are not real,
why do so many of us discuss them at such great length?
2. Individuation. Entities must be demarcated in some manner. In
general, we say they are spatiotemporally restricted and capable of re-
identification. Spatiotemporal demarcation is not always clear-cut, and
occasionally out-of-the-question (e.g., some entities in physics), yet the
predicates of theory still demand that such entities exist and even,
perhaps, specify their boundaries. Entities capable of individuation may
also be said to be discrete. Boundaries of all entities are sharp or fuzzy
depending upon the spatial and temporal scales used to describe them
(Hull 1976, pp. 185-186; Ghiselin 1981; Rieppel 1986, discusses the
problem of individuation of species in detail).
3. Irreducibility. Within the domain of a theory and the relationships
described by its predicates, entities should be irreducible. Irreducibility
implies that the entities will not be divisible into other entities that also fall
within the domain of the theory. The boundaries of entities are thus
theory-dependent (see also Hull 1974, p. 48). The notion of irreducibility
is especially relevant for species concepts because the usefulness of
biological theory hinges upon it. Given a particular definition of species,
irreducibility implies that entities meeting this definition cannot be sub-
334 JOEL CRACRAFT

divided into other, smaller entities which themselves play the role of
species within theories having them as their entities.
4. Comparability.The same kinds of entities must be recognizable and
comparable in order that the theory in question be empirically applicable.
Each theory specifies a series of predicates, and entities are comparable if
the same predicates apply to each of them (Gaukroger 1978, p. 39). If the
predicates and entities are inappropriately matched, then the entities
themselves may be improperly individuated or perhaps the theory may be
poorly or incorrectly formulated. Comparability is essential for a theory to
achieve empirical generality. We might have, for example, a theory of
planetary motion and wish to use it to explain the motions of the Earth,
Jupiter, the Milky Way, Pluto, and Alpha Centauri. That theory will likely
fail because the entities are not comparable - not all fall within the class
called planet. We either have to re-evaluate the entities or modify the
theory so that its predicates are applicable to each of the included entities.
The characterization of entities - the question of a correct ontology -
constitutes a major problem within evolutionary biology and nowhere is
this better illustrated than in the different ways species have been defined
and incorporated into theories about their origin. In the section that
follows, I will argue that the "synthetic theory of evolution" has incor-
porated very disparate concepts of species, so much so that it undermines
the very notion of a "synthesis." Then, in the section after that (VI), I will
show how the most influential species concept within evolutionary biology,
the biological species concept, is inadequate ontologically, thereby creat-
ing problems for any process-level theory utilizing biological species as its
entities. Finally, in section VII I will outline an alternative species concept
that avoids these ontological difficulties and which should be generally
applicable to theories about species origins for any group of organisms.

V. SPECIES CONCEPTS AND THE "EVOLUTIONARY SYNTHESIS":

THE PARADOX OF DISCRETE ENTITIES

The "evolutionary synthesis" is the loosely-held theory which states that

evolutionary change proceeds gradually as a result of genetic changes of
small phenotypic effect and as a result of the temporal ordering of genetic
variation by directional natural selection (Mayr 1980a; see also Eldredge
1985a). The "synthesis" is often said to be a unification of genetics,
systematics, and paleontology, the unification arising because all evolu-
tionary phenomena are considered explainable by (or consistent with)
those small genetic changes and the mechanism of natural selection.
There can be little question that many evolutionary biologists believe a
theoretical synthesis has been forged, for it is quite easy to find influential
evolutionists speaking of the theory of evolution or the synthetic theory
 SPECIES CONCEPTS AND THE ONTOLOGY OF EVOLUTION 335

(e.g., Charlesworth et al. 1982), as if there is a conceptually coherent body

of theoretical statements constituting a general, unified theory. But at the
very least, any unified theory requires a unified ontology, and from the
writings of proponents of the synthesis one can only conclude there exist
many different answers to the question, "What evolves?".
I will not undertake an extended discussion of the "units of evolution"
controversy but will address this issue from the standpoint of species
concepts. The argument here is that the use of widely disparate species
concepts within the component disciplines of the synthesis, along with
confusion over species within individual disciplines, undermines any claim
for the existence of a coherent theory of evolution.
The synthesis can be said to have arisen from three key documents
(Eldredge 1985a): Dobzhansky (1937), Mayr (1942), and Simpson
(1944). Dobzhansky and Mayr, because they saw species as reproductive
units within local communities, tended to have much more similar views
of species than either did with Simpson. Yet, neither had a consistent
ontology: both were unclear at times about what were the "units of
evolution," and over the years each expressed uncertainties about the
reality of species.
Even prior to writing "Genetics and the Origin of Species," Dobzhansky
(1935) apparently did not consider species to be the sole focal point of
evolutionary change: "the only biological category possessing an undisput-
able [sici ontological significance is that of a living individual" (1935: 344).
Despite this nominalistic viewpoint, he continues in this remarkable paper
to outline a concept of species that is widely used even today within
evolutionary biology:

Considered dynamically, the species represents that stage of evolutionary divergence, at

which the once actually or potentially interbreeding array of forms becomes segregated
into two or more separate arrays which are physiologically incapable of interbreeding.
The fundamental importance of this stage is ... that it is only the development of
the isolating mechanisms that makes possible the coexistence in the same geographic
area of different discrete groups of organisms. (Dobzhansky 1935: 354).

A key phrase of this quotation - "a species represents that stage of

evolutionary divergence" - signals a difficulty that many workers have
had in defining and conceptualizing species. Dobzhansky viewed taxa
within the context of a transformationist framework of evolution: "races,
species, genera, and families are nothing more than different degrees of
separation in the process of phylogenetic divergence" (1941, p. 369).
Given this view, what is the entity of evolutionary change? It would not
seem to be species for they are considered to be no different than any
other taxon except with respect to their degree of differentiation. Here
Dobzhansky seems to imply that populations are the units of change and
taxonomic rank is bestowed entirely as a result of a subjective assessment
336 JOEL CRACRAFT

of the amount of divergence (coupled, when circumstances permitted, to

an assessment of reproductive isolation). Dobzhansky had considerable
appreciation for systematics, and in most of his writings he viewed the
organisms of the world as being divisible into rather discrete entities -
species. But he was also a Darwinian transformationist and, like Darwin,
found it difficult to subscribe to an evolutionary viewpoint and at the same
time maintain the unity and discreteness of species. The ontological
problem present in Darwin's work was not solved by Dobzhansky's more
advanced view of species.
Mayr (1942) addressed many of the same problems as Dobzhansky
and did not reach fundamentally different conclusions, even though his
discussion of species and speciation was far more sophisticated. His
definition of species (1940, 1942, p. 120) was very similar to that of
Dobzhansky, although Mayr objected to the "dynamic" definition of
Dobzhansky in which species were seen as a stage of a process rather than
as the outcome of that process. Despite this objection, Mayr proceeded to
use much the same phraseology: "potentially interbreeding," "isolating
mechanisms," "stage in the evolutionary stream," and so on. More sig-
nificantly for this discussion, Mayr also possessed an uncertain ontology,
one apparently not that different from Darwin or Dobzhansky. The
paradox of discrete entities - that entities cannot be discrete and still
transform over time - is also present in Mayr's thought:

... we must make an attempt at a species definition. In doing this we are confronted by
the paradoxical incongruity of trying to establish a fixed stage in the evolutionary
stream. If there is evolution ... we should find all kinds of species - incipient species,
mature species, and incipient genera, as well as all intermediate conditions. To define
the middle stage of this series perfectly, so that every taxonomic unit can be certified
with confidence as to whether or not it is a species, is just as impossible as to define the
middle stage in the life of man ... (1942, p. 114) 3.

This quotation is not an isolated instance in Mayr's discussion. His

doubts about species reality are raised repeatedly in different guises. Some
are rather explicit:

The delimitation of species which do not belong to the same time level (allochronic
species) is difficult. In fact, it would be completely impossible if the fossil record were
complete ... The change Ibetween ancestors and descendants] is slight and gradual and
should, at least theoretically, not permit the delimitation of definite species. In practice,
. . .gaps in our knowledge make convenient gaps between the "species" (Mayr 1942, p.
153; italics added).

Dobzhansky and Mayr have no difficulty at all in subscribing to the

reality of species when analysis is confined to a single time plane or to
populations within local communities. Both recognize that delineating
species may be difficult at times, but the ontological question can be
answered: species are real entities and are the units of evolution. On the
 SPECIES CONCEPTS AND THE ONTOLOGY OF EVOLUTION 337

other hand, both Dobzhansky and Mayr abandoned the notion of reality
when species are projected back into time. In these cases, presumably,
populations are the "units" of evolution.
Mayr has attempted to solve this ontological dilemma by defining
species nondimensionally, that is, by restricting the use of the biological
species concept essentially to local communities where reproductive
interactions among differing populations can be observed. Some biologists
see this as a useful solution: "some workers are recognizing slowly that
evolutionary theory can be developed only on the nondimensional species
concept" (Bock 1981: 645). The relative merits of species definitions,
including the biological species concept, will be discussed in subsequent
sections, but it should be noted here that attempting to define away the
problem of species reality does not itself constitute a viable solution. For
one thing, Mayr's "solution" is actually an attempt to solve what is
perceived to be an epistemological, not an ontological, problem: namely,
how do we delineate species objectively in space and time? Even if the
"nondimensional" definition were to solve the epistemological problem,
and I shall argue it does not, the ontological issues remain: Are species
real, discrete entities, and are they the units of evolutionary theory?
Simpson's approach (1944) to evolution was distinctly different from
Dobzhansky's and Mayr's. Indeed, his entire language was different. He
was able to discuss speciation (1944, pp. 199-202), for example, and
mention the word "species" rarely and only in passing. Species concepts
and speciation played little stated role in his evolutionary epistemology.
Although his intellectual debt to Dobzhansky was clear, Simpson forged
his own paleontological view of the evolutionary process, albeit one
derived from the gradualistic worldview of traditional Darwinism. His was
an evolutionary theory in which populations and higher taxa seem as
important as species. For Simpson, populations are the primary units of
evolution. Simpson shared with Dobzhansky the belief that species and
genera (and even still higher taxa) are subjective designations we apply to
populations in order to signify their amount of divergent change.
Through virtually his entire career, Simpson saw species as arbitrary
segments of continuously changing lineages (1961, p. 165). To be sure he
acknowledged other conceptions of species, particularly that of the BSC,
but as a paleontologist and evolutionist he was reluctant to accept a
species concept that viewed species atemporally. This was brought out
forcefully in correspondence to Mayr:

I do not think the species problem has been solved during the synthesis, if by solution is
meant a definition and criteria generally acceptable to biologists and generally applic-
able without equivocation to all kinds of organisms. In that sense there simply is no
possible solution. The so-called biological definition has no time dimension and is
indeed nonevolutionary or preevolutionary, hence not really either an outcome of the
synthetic theory of evolution or integral to it. A truly evolutionary concept of species,
338 JOEL CRACRAFT

still open to various difficulties, has been reached by me and others and is integral with
the synthetic theory" (Simpson 1980, quoted in Mayr 1980, pp. 462-463).

This quotation by one of the acknowledged architects of the synthesis

underscores the thesis of this section, namely, that the synthesis could
not have encompassed a unified theory of evolution because it never
developed a coherent ontology. Proponents of the synthesis disagreed
over the identity of the units, or entities, that were said to evolve.
Dobzhansky and Mayr generally identified species as units even though
they often saw species as arbitrarily delimited and of questionable reality.
Simpson was more straight-forwardly sceptical of species reality. All three
came closest to agreement when specifying populations as things that
evolve, but throughout most of their work neither Dobzhansky nor Mayr
saw the fundamental divisions of nature to be populations 4 . Their view of
the world, shared by most biologists, is that species are those fundamental
units. But widespread agreement on this point still does not resolve the
difficulties for evolutionary theory, because many different species con-
cepts have been proposed and each has its own ontological and epistemo-
logical implications for any proposed theory of evolution.

VI. THE BIOLOGICAL SPECIES CONCEPT AND EVOLUTIONARY THEORY

The biological species concept (BSC) is widely accepted within evolu-

tionary biology (e.g., Bush 1975, White 1978, Futuyma 1986). By virtue
of accepting that definition, however, evolutionary biologists establish a
specific epistemological approach toward evolutionary theory. Species are
reproductively disjunct, or isolated, units; speciation is the origin of repro-
ductively isolated units; the genetics of speciation is that genome change
which causes reproductive isolation; reproductive isolation is the primary
consequence of the evolutionary process.
Assume at the outset that there exist real units in nature, units that are
produced by some unspecified process of descent with modification. We
could ask how those units might be delimited and develop criteria for
sorting individual organisms into their appropriate unit. But different
processes might be expected to produce different units, and how we
understand those units to be constituted could influence the way in which
we describe these processes. According to proponents of the BSC, nature
can be partitioned into units on the basis of reproductive isolation, yet as
many critics of the BSC argue, reproductive isolation is not the only
criterion that might be used to recognize the fundamental entities we call
species. Resolution of this conflict resides in establishing which definition
best permits us to understand the patterns and processes associated with
the temporal and spatial history of the units themselves. This requires that
 SPECIES CONCEPTS AND THE ONTOLOGY OF EVOLUTION 339

we simultaneously develop a theory about descent with modification

which incorporates these units, but units based on different definitions are
not necessarily equivalent.
Criticisms of the BSC have taken many forms (e.g., Ehrlich 1961, Sokal
and Crovello 1970, Cronquist 1978, Rosen 1978, 1979, Levin 1979,
Raven 1980, Cracraft 1983, Rosenberg 1985). This discussion empha-
sizes certain problems of the BSC that arise from a consideration of the
ontological requirements of evolutionary theory. Among other things, it
will be argued that the ontology implied by the BSC is incorrect: the focus
of evolutionary theory should not be concerned with reproductively
isolated units but with differentiated units and this should be reflected in
its ontological structure.
We begin with two criticisms, both of which are readily acknowledged
by proponents of the BSC at the same time their importance is ignored.
First, the BSC lacks generality inasmuch as it cannot be applied to asexual
organisms (Mayr 1963, pp. 27-29; Rosenberg 1985, pp. 192-194). The
point of interest here is not that a species concept (even the BSC) might
be difficult to apply in practice (Mayr 1963, p. 29); rather it is that the
BSC, in principle. cannot be applied to an important component of
biological diversity. The logical consequences of this are several. Given
that biological species are the correct currency of evolution, we might
conclude a general theory of evolution, applicable to both sexual and
asexual organisms, cannot be specified. Instead, we may need to recognize
two (or more) different theories, each with their own distinct entities. On
the other hand, perhaps a general theory does exist, in which case we
would have to conclude that biological species do not represent the
relevant evolutionary units for this theory.
Is there a way, short of ignoring the above-noted problem altogether (as
many biologists have done for the past 50 years), to salvage the BSC? One
solution is to redefine the BSC such as Mayr (1982, pp. 273-275) has
proposed: "A species is a reproductive community of populations (repro-
ductively isolated from others) that occupies a specific niche in nature."
Reproductive isolation fails as a criterion for asexual species, so one
therefore invokes differences in niche occupancy (Mayr 1982, p. 275).
Yet, this newer version of the "biological species concept" merely in-
creases our difficulties for it seems certain that "niche" cannot be objec-
tively defined in the absence of knowledge about the identity and ecology
of entities occupying that physical-biological "thing" called the "niche."
Species must be recognized prior to discriminating niches. As a con-
sequence, this redefinition of the BSC borders on circularity (see also
Rosenberg 1985, p. 194). Ghiselin (1987) views an attempt to define
species ecologically (see also Van Valen 1976) as "an act of desperation,"
but, ironically, in Mayr's case it is an act designed to save the BSC, which
Ghiselin supports so strongly.
340 JOEL CRACRAFT

A second major difficulty of the BSC was noted in the previous section,
namely its temporal inextensibility. Simpson (in Mayr 1980, pp. 462-
463), recall, believed the BSC to be "nonevolutionary" because of its
"nondimensional" character. Advocates of the BSC try to imply this
limitation is insignificant:
"the species concept has its full meaning only where populations belonging to different
species come into contact. This takes place in local situations without the dimensions of
space (geography) and time . .. The function of the species concept is to determine the
status of co-existing individualsandpopulations"(Mayr 1982, p. 292; italics added).

The temporal dimension is not the friend of the biological species

concept:
"It is only in the cases of a sequence of ancestor-descendant species which transform
gradually into each other in a single phyletic lineage that a sharp delimitation between
temporal species taxa is impossible. Here biological evolution fails to accomodate the
wishes of the taxonomist. Fortunately, the fossil record is more accomodating. Its
deficiencies usually provide sufficient gaps in the lineages to permit a delimitation of
vertical species taxa, as artificial as this may be. It seems that we will have to accept this
compromise solution ... " (Mayr 1982, p. 295).

This view of species ontology and its role in evolutionary theory and
analysis is, to say the least, remarkable. Mayr recognizes (e.g., 1982,
p. 286) that evolutionary analysis demands temporal extensibility, but
then claims species can only be delimited artificially when so extended. If
that argument is accepted, it implies evolutionary theory, which certainly
is about patterns and processes extending over time, cannot be about
"species" if the entities said to be species are little more than artifacts of
the fossil record.
What leads some biologists and paleontologists to believe that species
"transform gradually into each other in a single phyletic lineage"? One
obvious answer is that the belief is a consequence of how we define
species in the first place. "Empirical" observations such as "species trans-
formations" are interpretable only within the context of prior conceptuali-
zations about the nature of species. If species are defined so as to encom-
pass lineage segments between speciation events or branch-points (Wiley
1978; Cracraft 1983), then by definition sequential species within those
lineages would simply be artifacts and without any ontological status, thus
eliminating any need for gaps in the fossil record to be used to individuate
species. Defining, or conceptualizing, species in terms of branch-points
provides an objective basis for individuating species, but as with any
definition we can never be assured of being free of empirical difficulties
due to inadequacies in available data. Assigning species-rank to a group of
individual organisms is a scientific hypothesis (Eldredge and Cracraft
1980), but such hypotheses should not be proposed about unreal or
artifactual entities. To talk about species when those species are arbi-
 SPECIES CONCEPTS AND THE ONTOLOGY OF EVOLUTION 341

trarily delimited, constitutes "merely a statement reflecting the mental state

of the investigator" (Brothers 1985).
Surprisingly, failures to accomodate asexual species or situations span-
ning evolutionary time may not be the most significant difficulties for the
BSC. A more general problem exists: the BSC, because it depends upon
reproductive disjunction as its criterion of individuation, systematically
misidentifies the "units of evolution" and thus confounds evolutionary
analysis. This judgment of the BSC is best explored by introducing an
alternative concept of species.

VII. THE PHYLOGENETIC SPECIES CONCEPT: NATURE DIVIDED AT ITS

JOINTS

In arguing against a species concept applicable across all organisms, Mayr

(1987, pp. 37-38) draws the only logical conclusion from a committment
to the BSC: the history of life cannot be united within a single explanatory
framework. Many biologists and philosophers of biology surely will find
this position difficult to accept 5 . One step toward solving this dilemma is
fairly obvious: abandon the biological species concept, and in so doing
some critical ontological difficulties of the synthesis will be eliminated
immediately. Inasmuch as biological species do not carve nature at its
joints (Rosenberg 1985, p. 197), biology is faced with formulating an
alternative species definition. Although this has been a persistently trouble-
some task, the previous discussion suggests a deceptively simple solution:
equate species with evolutionary units. Accordingly, a species can be
defined as an irreducible cluster of organisms, within which there is a
parental pattern of ancestry and descent, and which is diagnosably distinct
from other such clusters. Species are thus basal, differentiated taxa
(Nelson and Platnick 1981; Cracraft 1983).
This definition is no less "biological" than the BSC because diagnos-
ability could be based on any intrinsic attribute. The definition manifests
what we expect from our contemporary understanding of the evolutionary
process: ancestral taxa differentiate and produce new, diagnosably distinct,
descendant taxa. It is important to address the issue of species individua-
tion or recognition 6, for some biologists or philosophers might be tempted
to label the phylogenetic species concept "typological" merely because it
places considerable reliance upon the use of diagnostic characters. The
definition, however, also has the phrase "parental pattern of ancestry and
descent" (see also Nelson and Platnick 1980, p. 12; Cracraft 1983), and I
suggest that most biologists use pretty much the same methods (but
perhaps interpret the results differently) when individuating basal taxa
(here termed species). By and large two criteria must be applied: (1) one
or more diagnostic characters (of any kind as long as they are heritable),
342 JOEL CRACRAFT

and (2) reproductive cohesion (not disjunction). Both are often necessary
because the diagnostic characters of a species may be restricted to a single
sex, ontogenetic stage, or morph; observations on reproductive cohesion
of populations help us to resolve such difficulties. But knowledge about
diagnostic characters will always be essential. Even in cases of sympatry
and hybridization of two evolutionary units, we know there are two units
simply because (1) each is different from one another, and (2) each type is
observed to be reproductively cohesive outside the zone of sympatry. The
criterion of reproductive disjunction, on the other hand, is clearly of
secondary importance for without the presence of some diagnostic charac-
ter, how could we ever recognize that two taxa were hybridizing or that
two "sibling species" were not hybridizing? Without those characters we
would see only a single, reproductively cohesive population. In practice,
reproductive cohesion is a component of all species concepts including
purely "morphological species concepts" (proponents of such definitions,
after all, do not place sexes in different species except through uninten-
tional error). Reproductive disjunction, on the other hand, is a criterion
only of the BSC, and because of this many advocates of that definition
often imply their's is the only one to use reproductive criteria. In speaking
of the criteria used to delimit "individuals," both Hull and Ghiselin have
stressed the cohesiveness of such units. When Mayr speaks of these
individuals (e.g., 1987, p. 19), an altogether different view emerges: "No
one questions the fact that organisms which belong to a species are united
by the joint possession of a set of isolating mechanisms .... " This "fact"
can be questioned, however, for "joint possession of a set of isolating
mechanisms" cannot by itself delineate a species. "Isolating mechanisms"
are fortuitous effects of differentiation and in the majority of cases merely
constitute the absence of a recognition system present in another species
(Paterson 1985). Relative to any particular species, many other species
"share" the absence of some recognition mechanism and hence are "jointly
isolated" from that species. Diagnostic characters and reproductive cohe-
sion, not reproductive disjunction, reveal the boundaries; without the
former, the analysis of reproductive disjunction is impossible. It follows
that the use of reproductive cohesion in discussions of species individ-
uality does not require loyalty to the BSC. So-called "biological species"
are not comparable entities because some include many differentiated
evolutionary units, whereas others include only one 7 . This disparity results
from subjective assessments of potential reproductive disjunction. When
units are reducible and noncomparable, as biological species often are,
evolutionary analysis can be seriously confounded (Rosen 1978, 1979;
Cracraft 1983).
Space does not permit a detailed explication of the phylogenetic con-
cept of species (see especially Cracraft 1983), and this is not the forum to
engage in an extended scientific debate over alternative species concepts.
 SPECIES CONCEPTS AND THE ONTOLOGY OF EVOLUTION 343

Yet, several philosophical issues are raised when comparing the BSC with
the phylogenetic species concept. Biological and phylogenetic species are
equivalent entities only when the former are monotypic. If monotypic and
"polytypic" biological species are not comparable entities - in many
respects polytypic species comprised of two or more discrete, basal taxa
are analogous to genera - then what does it mean to claim "biological
species are individuals"? Are all species, no matter how defined, individ-
uals? Certainly any of those species can be given a proper name - need it
be pointed out that Homo sapiens is a "polytypic species"? - but this does
not imply those "species" will necessarily exhibit characteristics of individ-
uals such as spatial or temporal continuity. And in what sense can the taxa
placed in these different species be said to constitute "real entities"? Some
surely are, but the extent to which other species are taxonomic conven-
iences (or artifacts), as is perhaps the case when differentiated allopatric
populations are placed in the same biological species because it is thought
they might be capable of hybridization, we are faced with ontological
questions that have not yet been explored in sufficient depth.
Biology has not yet solved the species problem, and the scientific issues
will continue to be debated. If philosophers are to contribute to its
solution their participation will necessarily require commentary on scien-
tific issues. Despite strong sympathies toward the viewpoint of "species as
individuals," I question whether this philosophical concept will have
anything more than an indirect bearing upon the scientific debate over
species concepts. The criteria used to recognize individuals - spatio-
temporal restriction, part-whole relationship, spatiotemporal continuity or
cohesion (Hull 1976, 1980; Ghiselin 1981; Eldredge 1985b) - would
seem applicable to nearly all kinds of species, whether they be biological,
phylogenetic, evolutionary, or ecological, and arguments to that effect
can be found in the literature. The usefulness of individuality, instead,
resides in forcing biology to address the ontology of evolutionary theory.
If that theory requires species (of whatever kind) to be individuals, then
certain approaches to species will have to be discontinued. Species can no
longer be treated as arbitrarily defined conveniences through space and
time. Such a view will have broad implications for both neontology and
paleontology.

NOTES

* I want to thank David Hull and Niles Eldredge for their comments on this manuscript. I
am also grateful to the National Science Foundation (through grant BSR-8520005) for
support of this research.
These biologists might argue, perhaps, that the "species as individuals" controversy is a
philosophical question and cannot be decided on empirical grounds. Even if true, this
344 JOEL CRACRAFT

would not vitiate its importance for the conceptual organization of biological theory, which
certainly could have empirical consequences. Individuals are supposed to be discrete
entities; if each species is a discrete entity, then species are individuals, not classes; if
species are not discrete entities, then there would seem to be little reason to talk about
them in the ways that we do. Some philosophers (e.g., Kitts and Kitts, 1979; Caplan 1981)
see particular species as being classes or natural kinds, yet because these species are
spatiotemporally restricted (at least most biologists would say they are), it is doubtful that
they could be the subject of laws (Hull 1981 b).
2 It is important to keep in mind, however, that one should not equate the "unit of

evolution" with the "unit of selection" (e.g., Caplan 1981). Many biologists accept that
species "evolve" while at the same time recognizing that selective processes act on lower-
level entities such as genes or individual organisms.
I As we shall see, Mayr draws an incorrect analogy. We should not compare the middle
stage of a man's life to his whole life, but an individual's life to the history of a population.
A man's middle life may be delimited subjectively, but most would consider a man's life as
being rather well defined. Hence, a person is an individual whereas a "middle life" is not.
4 Apparently Mayr (1987) has moved toward equating "population" and "species," which
is a curious position for someone advocating the BSC.
s Thus, I share Ghiselin's (1987) perspective that, in principle, we want to seek an
evolutionary theory which is as generalizable as theories in the physical sciences. I agree
also that the individuality of species is an important philosophical ingredient of that goal. I
disagree, however, that the BSC provides us with the proper ontology to obtain that goal.
In fact, proponents of the BSC have already admitted that it fails in that regard. Ghiselin
(p. 15) too easily dismisses criticisms of the BSC. "The biological species definition," he
says, "turns out to be fully applicable to those plants in which species actually do exist." It
is true that the BSC seems to apply when reproductive isolation, the cornerstone of the
BSC, is observed. But there is a logical and biological problem here: Ghiselin seems to
imply that if the criterion of reproductive isolation cannot be applied, then species might
not exist (I am not arguing here that this might not be a defensible position). An alternative
interpretation is that species actually exist in these groups, but the BSC is unable to delimit
them. It fails, not only in asexual groups, but in many taxa in which reproductive isolation
is not always an outcome of evolutionary divergence. Like Ghiselin, however, I reject calls
for "pluralism" in species concepts (e.g., Scudder 1974; Mishler and Donoghue 1982;
Kitcher 1984; and now Mayr 1987) on both philosophical and scientific grounds.
' Here and elsewhere I use the notions of individuation, recognition, delimitation, and
diagnosability as rough equivalents. I am concerned with the empirical evidence we use to
identify a collection of individual organisms as a discrete taxon we call a species.
7 Many "biological species" (perhaps the majority) contain a variable number, sometimes
quite large, of "subspecies." In some instances these subspecies represent truly differen-
tiated taxa, whereas others are "arbitrary" subdivisions of continuous spatial variation. In
many cases subspecies are geographically isolated from each other and their placement in
the same "biological species" is almost always based on an assessment of relative phenetic
(overall) similarity. Close similarity is assumed to mean they would interbreed if they were
to eventually come into contact, and a greater degree of phenetic dissimilarity is taken to
mean they would not interbreed if they were to become sympatric. These practices of
"evolutionary systematics" (Mayr 1942, 1963) clearly imply considerable subjectivity in
"individuating" evolutionarily important entities. Using this systematic philosophy and
procedure, "biological species" cannot possibly be comparable among themselves because
they often do not represent the same "thing."
 SPECIES CONCEPTS AND THE ONTOLOGY OF EVOLUTION 345

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