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A Radical Solution to the Species Problem

Michael T. Ghiselin

Systematic Zoology, Vol. 23, No. 4. (Dec., 1974), pp. 536-544.

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A RADICAL SOLUTION TO THE SPECIES PROBLEM

Abstract
Ghiselin, M. T . (Department of Zoology and Bodega Marine Laboratory, University of
California, Berkeley, P. 0. Box 247, Bodega Bay, California 94923). 1975. A radical solution
to the species problem. Syst. 2001.23:536-544.-Traditionally, species (like other taxa)
have been treated as classes (universals). In fact they may be considered individuals (par-
ticular things). The logical term "individual" has been confused with a biological synonym
for "organism." If species are individuals, then: 1) their names are proper, 2 ) there cannot
be instances of them, 3 ) they do not have defining properties (intensions), 4 ) their con-
stituent organisms are parts, not members. "Species" may be defined as the most extensive
units i n the natural economy such that reproductive competition occurs among their parts.
Species are to evolutionary theory as firms are to economic theory: this analogy resolves
many issues, such as the problems of "reality" and the ontological status of nomenclatorial
types. [Biological species; systematics; philosophy.]

It would appear that the philosophy of uals. This paper, however, only aims to
taxonomy is about to undergo a major show that such a position has attractive
upheaval. Symptomatic is its Gordian knot, qualities from the point of view of logic
the species problem. Some years ago and biology alike, and that it is perhaps not
(Ghiselin, 1966a) I attempted to cut it with so radical as one might think. The basic
the sword, casually remarking that, in the point (Ghiselin, 1969, 1974) is that multi-
logical sense, species are individuals, not plicity does not suffice to render an entity
classes. So entrenched had the habit be- a mere class. In logic, "individual" is not
come of treating species as universals rather a synonym for "organism." Rather, it means
than as particular things, and their names a particular thing. It can designate systems
as general rather than proper, that it took at various levels of integration. A human
some time before this pronouncement was being is an individual in spite of being
taken seriously. The position in question is made up of atoms, molecules and cells. An
hardly new. It goes back at least to Buffon, individual, furthermore, need not be phys-
but I believe this was the first explicit use ically continuous. The United States of
of it against what has been misleadingly America is an individual in the class of
called the nominalistic species concept. national states, and this is true in spite of
Even Ruse ( 1973), a philosopher who cites the interposition of Canadian territory and
the foregoing work, passes the notion over international waters between Alaska and
in silence. On the other hand Hull (1974) the rest. (For a philosophical discussion of
has lately endorsed the idea that, from the individuals see the book on that subject by
point of view of evolutionary theory, bio- Strawson, 1959.)
logical species and monophyletic taxa are It is characteristic of individuals that
individuals. And Mayr (1969a), while not there cannot be instances of them. Thus,
going so far, strongly emphasizes the point the class of national states has instances:
that species are more than just nominal the United States of America is a national
state. We would not, however, say, "Cali-
classes. It would appear that the subject
fornia is a United States of America." This
deserves further analysis and comment. is true in spite of the fact that we may treat
any number of parts of the Union as a class,
PHILOSOPHY
as when we say that these "states" are
There are both advantages and dis- sovereign. California is a part of the United
advantages to treating species as individ- States of America.
SPECLES PROBLEM

Equally significant is that individuals do as particular species cannot have defining


not have "intensions." That is to say, there properties, the question is closed.
are no properties necessary and sufficient
to define their names. Of course, people, BIOLOGY
national states and species do have prop- The most significant question for biol-
erties, but these are not implied when we ogists is "Individual whats?" One answer
use the name. Much of the species problem is individual populations-in the sense of
has been the result of equivocal uses of genetical populations ( syngamea) . These
species names as universal and proper. are breeding communities-composite
Thus, some biologists will never say "John wholes-not to be confused with classes of
Smith is a Homo sapiens," but insist that organisms sharing certain intrinsic genetical
correct usage demands "John Smith is a properties. Of course one has to add some-
specimen of Homo sapiens." In other thing, namely that they are reproductively
words, some consider the taxonomic term isolated, to differentiate "species" from
to designate a class of people ("all men") "subspecies" and other populational cate-
others as a single thing ("humanity" or gories. This, the biological species defini-
"the human race"). tion, is adequate, but it is still affected by
Some confusion, and indeed some con- certain problems of interpretation and im-
tinuing uncertainty upon the part of logi- plication which suggest that a better way
cians, results from the fact that proper might be found to define this category.
names can imply or suggest certain prop- Some definitions, albeit technically sound,
erties. It does not seem to me that this could use some improvement. "Gold," for
means that proper names really do have example, could be unequivocally defined
intensions, but perhaps some would dis- as the element with an atomic number of
agree. A descriptive epithet facilitates 79, but for most of us, this is not very
communication and aids the memory. Con- informative.
sider, for example, Phallus impudicus Lin- One solution is to try to define the species
naeus, 1753. But let us not confound poetry and other categories in terms of the causes
with logic. Whether a name be apt or a of evolution. This is one of the main reasons
misnomer is not the same question as what why the usual formulation of the biological
properties a thing must of logical necessity species definition is so attractive. Gene
possess if the name is to be used. flow and reproductive isolation obviously
As is well known, much confusion is profoundly influence the properties of
avoided in dealing with such matters when organisms. But might not something else
one clearly distinguishes between categories be more important? One possibility is com-
and taxa. Taxonomic categories, in the petition, including natural and sexual selec-
sense of all taxa of a given rank, are like tion. El~ewhere (Ghiselin, 1974) I have
"national state": everybody agrees that argued at some length, giving historical
these entities are classes. The point at issue documentation, that biologists have mis-
is whether taxa are likewise classes, or interpreted the notion of competition and
whether, like California, they are individ- failed to appreciate its full significance.
uals. It may be that some taxa (e.g. Mam- Ecological data have, to be sure, entered
malia) are classes, whereas particular into the thinking of systematists, but no-
species are individuals. In traditional anal- body seems to have considered the pos-
yses of the species problem, which presup- sibility of defining categories in terms of
pose that all taxa are classes, there has been modes of competition. Let us consider one
some confusion over whether the names of possibility.
categories should be defined so as to specify One can hardly deny that different kinds
the defining properties of taxa. In the light of competition occur between species and
of the present analysis, one would say that within them. Interspecifically we have the
SYSTEMATIC ZOOLOGY

struggle for the means of existence only. This may be dismissed as merely a version
Intraspecifically there occurs a competition of a paradox invented by Heraclitus (frag-
with respect to genetical resources as well, ment LXXXI in the Loeb Classical Library
and even the resources contended for by edition) : one cannot step in the same river
organisms irrespective of species in the final twice. Who but a madman or a philosopher
analysis are directed toward the intra- would maintain that he himself does not
specific struggle for reproductive success. exist because he changes? Are we all fig-
Species, then, are the most extensive units ments of our own imaginations?
i n the natural economy such that reproduc- The rather forced dimensional-nondimen-
tive competition occurs among their parts. sional dichotomy emphasized by Mayr
I t would be circular, if perhaps not vi- (1963) is now seen to be irrelevant. Species
ciously so, to say that species and lower need not be defined solely in relation to
taxa are those entities within which intra- other species. "Species" is not a relational
specific competition occurs. Some might concept like "brother," but something com-
object to the use of "reproductive" in the parable to "man." A firm is a firm because
foregoing definition, but there seems to be it forms a closed system of a given kind.
no better alternative. By "reproductive I t can compete with craftsmen and firms
coinpetition" is meant a competition with outside itself, and is characterized by a
respect to genetical resources as such, not particular kind of internal organization. An
just any competition involving reproduc- only child cannot be a brother, but an
tion. economy might contain but a single firm.
Species are to evolutionary theory as The difficulties surrounding "potential
firms are to economic theory. And organ- interbreeding" become a dead issue too. If
isms are productive units in both, compet- a species is an individual it hardly matters
ing in different ways within their larger whether it is interbreeding at any given
units and with the parts of comparable instant. All the "members" of a species are
units. Likewise some organisms exist au- competing reproductively with all the
tonomously-as asexual creatures and self- others, irrespective of the distance between
employed craftsmen. This analogy has its them or the existence of temporary spatial
limitations, of course, not the least being discontinuities between their component
that firms can reasonably be said to be populations. Intraspecific competition is
adapted, while species cannot. Nonetheless more perfect between organisms in close
the analogy can be shown to help resolve proximity, but the whole gene pool is none-
the paradoxes and conceptual difficulties theless at issue. One should realize that
that have so long plagued the subject. all organisms, everywhere, like all the
Species are individuals, and they are real. productive units in the world-wide political
They are as real as American Motors, economy, are able to compete with one
Chrysler, Ford and General Motors. If it another to some degree. All can do that
be true that only individuals compete, then which diverts resources from the rest to
species as well as organisms can compete, themselves. They do so more or less in-
just as corporations and craftsmen can. But tensely, or directly, or perfectly, and in
clearly there exists a profound difference diverse modes, but compete they do, and
between two species competing, and com- they can compete at a distance.
petition between organisms belonging to Some theorists have expressed the view
different species. that the existence of "different kinds" of
Burma (1949) claimed that because species creates a problem for the biological
species differ from moment to moment in species definition. Some have felt that there
geological history, they are, therefore, not should be special categories (or kinds of
the same, and hence that they are mere categories) for designating these. The
concepts without real existence in nature. analogy with economics is particularly use-
SPECIES PROBLEM

ful here. Differences in the range of diver- exist side by side with various stages in
gence within species-as when some are the formation and dissolution of trusts.
monotypic, others polytypic-create no The morphological, genetical and physio-
problem. After all, some corporations are logical species concepts suffer from a com-
like Mitsubishi, others like the local fish mon fault. They treat composite wholes as
market, but both can be corporations. I t if they were classes defined in terms of the
has been most unfortunate in this context intrinsic properties of their members. If
that niche theory has been so intimately one were to define "craftsman" in terms of
bound up with the notion of species as the properties of cells in certain organisms,
classes, so that niches are conceived of as it might seem odd. Likewise, we are not
"abstractly inherited attributes of sets of interested in the properties of employees
like organisms. Hence a typological concep- when we define "firm," but rather in the
tion of the niche could scarcely be avoided. properties common to firms generally. The
It becomes easier to reason about such attributes of organisms are not defining of
matters when one realizes that a craftsman the names of social groups, in spite of the
or other organism makes his living in a fact that social groups must have constitu-
sense distinct from that of operating within ent organisms. That John Doe has a par-
a sector or range of sectors of an economy. ticular set of genes is about as relevant to
The economic activity of both can vary his being a specimen of Homo sapiens L.
from time to time and place to place. as it is to his working for the manufacturers
No paradox arises when a single factory of Brand X. Hence I cannot agree with
constitutes the entirety of a firm. Likewise, Mayr (1969a:29) that the species "receives
some biologists wish to erect "agamo- its reality from the historically evolved,
species" for species-less organisms. I t seems shared information content of its gene
rather like creating imaginary firms for the pool." The species and the firm are "real"
self-employed. And no economist would in the sense that they designate entities
ever erect "chronofirms" for the same firm which exist: particular species and firms.
at different times, although the name of a And organisms are no more vessels for
firm might be changed when it becomes genes than they are vessels for industrial
reorganized. skills. Geneticists on the one hand, and
Consider, too, the objection of Ehrlich employers on the other, may view them in
and Raven (1969) to the biological species such light; but the disadvantages to doing
concept on the grounds that it overem- so should be obvious.
phasizes integration and gene flow within One problem with the "phenetic species
species. Whether this be a valid criticism concept" (see especially Sokal and Cro-
or not, it is circumvented by shifting the vello, 1970; Sneath and Sokal, 1973) is that
emphasis from integration and gene flow it treats species as classes defined in terms
to the mode of competition. Firms are firms of the traits of organisms, rather than as
irrespectively of how tightly organized and individuals having the properties necessary
closely integrated they may be, and they and sufficient for membership in the species
remain basic units of organization in spite category. The pheneticists' position is that
of the fact that local productive units are we should put things together according to
important too. That limited exchange of their degree of resemblance. Yet, from the
genes may occur between species creates point of view of evolutionary theory, it is
no more difficulty for the species concept the causal nexus that matters. Of course,
than does the fact that someone might work erecting classes on the basis of shared traits
for two firms create difficulties for the has its place, but this is not the issue. Some
notion of a firm. Gradual development of taxonomists may not care whether two
reproductive barriers creates intermediate organisms belong to different (cryptic)
conditions, but closed economic systems species, but it certainly matters to them.
SYSTEMATIC ZOOLOGY

All banks are not the same when we are less, subspecies may be expected to each
depositing money, no matter how much have a characteristic local mode of com-
they look alike to us. petition, and what has been called their
The realization that species are individ- "reality" may be compared to that of any
uals helps us to understand some of the local productive unit. Yet as I have pointed
important daily activities of taxonomists. out (Ghiselin, 196613) some lower categories
How, for example, do they define the names do not refer to populations: the morph, the
of species? If species were universals, then forma sexualis, the monstrosity, etc. These
their names would have defining charac- are strictly nominal classes, and for that
ters. The problem of there being much reason are best left out of the hierarchy.
diversity within certain species has been Are the taxa ranked at categorical levels
solved in a rather makeshift fashion by higher than the species merely conceptual?
considering them to be "disjunctively" or The economic analogy here seems rather
"polythetically" defined-to view them as remote. Is the automobile industry real?
"clusters" of similar objects, but without One's answer depends upon his meta-
any one combination of properties being physics, but at least industries have not the
necessary and sufficient to qualify an entity same ontological status as firms. Whatever
for membership. But according to the view answer is best, it may be worth while to
being examined here, species names are explore the possibility of treating higher
proper names-like American Motors. I t is taxa as sectors of the natural economy.
not only difficult, but logically impossible, Inger (1958) suggests some thoughts along
to list the attributes necessary and sufficient these lines for the genus, as have Gisin
to define their names. None such exist, and (1960) and Whittaker (1969) for the higher
the only way to define these names is by categories. On the other hand we should
an ostensive definition. That is to say, by bear in mind the momentous difference
"pointing" to the entity which bears the between the natural and the political econ-
name. Much confusion about the role of omy, in that a phylogenetic nexus is the
nomenclatorial types has arisen because inevitable consequence of the laws govern-
they are used in a peculiar kind of ostensive ing reproductive communities. Firms are
definition. Attention is drawn to only a established out of heterogeneous elements,
part or aspect of the individual the name ones not necessarily arising out of a com-
of which is being defined. This entity, the mon source. Be this as it may, the economy
type specimen, is misinterpreted as an ex- of nature is competitive from beginning to
ample of a class, which it is not. Likewise, end, and assimilating so basic a process as
the diagnosis, which in fact only "purports competition into evolutionary classification
to differentiate the taxon," is often confused might greatly strengthen the conceptual
with a definition, which actually would foundations of taxonomy,
differentiate it. I t is as if one were to
define "Mobil Oil Corporation" by pointing HISTORY
to a single service station. One might wonder why all of this has
The perplexing issues of the defining not been self-evident for years on end. To
properties and ontological status of the give a full answer would require an exten-
categories lower than the species may be sive historical study, but the following
clarified in a similar vein. Subspecies are suggestions may help.
like local branches of a widespread in- Elsewhere (Ghiselin, 1972) I have pre-
dustrial enterprise, but the degree of dif- sented a theoretical scheme which attempts
ferentiation can hardly be so distinct as it to explain scientific innovation. For the
is at the species level, for there can be all present discussion, three main features of
sorts and degrees of closure, and closure is this scheme need to be singled out. In the
not the same as differentiation. Nonethe- first place, dealing with a problem depends
SPECIES PROBLEM

upon having an appropriate way of think- form a family" he should have realized that
ing about it. Method determines success. whatever it may be that they form, it is
Second, an ability to invent or discover a anything but a family. Poker does not
new way of thinking largely is contingent copulate with Canasta or mourn for the
upon some kind of input from without. This passing of Whist. Of course, Wittgenstein
external input gives us the third point: wasn't talking about that; but this is just
discovery often involves interdisciplinary the point: he should have.
transfer. But this transfer of concepts and I t seems curious that even Kant (1776,
ways of thinking may be non-existent, 1785), who made outstanding contributions
shallow, or profound, according to the to both science and philosophy, commented
habits and training of the scientists in- upon species and the views of Buffon, but
volved. For various reasons some fields of contributed nothing of significance to the
research and their practitioners are rela- philosophy of taxonomy. Yet one must
tively open, others relatively closed, to the remember that neither Buffon nor Kant
rest. It is partly a matter of personal situa- understood the competitive interactions
tion, and partly a matter of innovative that go on between species, between parts
mentality that determines how a given of different species, and within them. They,
scientist will behave in such respects. like all the pre-Darwinian biologists had a
The species problem has to do with teleological conception of the universe,
biology, but it is fundamentally a philo- featuring an harmonious and cooperative
sophical problem-a matter for the "theory natural economy here on earth. Their
of universals." There are, so to speak, world-view prevented biologists and philos-
"different kinds of groups," and someone ophers alike from thinking about species in
trained in logic should, one might think, terms other than "similarity and filiation"
long ago have stepped in and cleared up so that these groups almost had to be classes
the confusion. Such is demonstrably not -analogous to defining "the Smith family"
the case, for logicians ever since Aristotle, as that which resembles John Smith. No
and even quite recently, have treated spe- amount of study from this point of view
cies as classes (e.g., Gregg, 1950; Buck and would lead one to think in terms of natural
Hull, 1966; Lehman, 1967; Ruse, 1969, selection, a process involving interactions
1971). There may be quite a variety of between the organisms making up the
reasons for this practice, but one significant group.
point is that philosophers habitually seek Only when Darwin, a geologist, tran-
only to erect internally consistent schemes. scended the limitations of the traditional
They have little to gain by attempting to way of thinking was the problem of specific
discover how the living world really is transmutation possible. He reached beyond
organized, or how biologists in fact con- the limits of biology, to economics, and
ceive of it. Hence although there has been realized that evolution results from a repro-
much philosophizing about science, there ductive competition between individual
has been little that truly deserves to be organisms. Hence the Darwinian revolution
called the philosophy of science. The usual should be viewed as an ecological, and not
notions of logicians are adequate enough as a taxonomic, revolution, in the sense that
that these can be applied to scientific clas- a new ecology implied a new systematics,
sification without breaking down. Some- not vice versa.
thing special was needed if logic were to Yet Darwin's own outlook had significant
cope with biology as it really is. This limitations. He was so concerned with the
situation is unfortunate, for not only might transmutation of species that he gave insuf-
philosophers have much to contribute, but ficient emphasis to their origin. This led
they might learn something as well. Thus him to conceive of species more or less as
when Wittgenstein (1953:32) wrote "'games' open systems, rather than as closed systems.
SYSTEMATIC ZOOLOGY

In his later years, he to some extent ad- classes with members (Plate, 1914; Turrill,
mitted this, and agreed that isolation, albeit 1942; Gregg, 1950). A few have maintained
not necessary for evolutionary change, is a kind of idealism: to Agassiz (1857) spe-
helpful in speciation (letters in F. Darwin, cies were thoughts of the Creator, which
1887: Vol. 111, pp. 157-162). Confusion are real. Many biologists have conceived
between these two processes continues to of species as individuals, but instead of
this day, especially in the writings of his- putting it in these terms, they have merely
torians (see Ghiselin, 1973). The problem said that they are not the same kind of
becomes somewhat less difficult when one groups as classes are. Some have viewed
treats both species and organisms as indi- them as phylogenetic units (e.g., Harper,
viduals. 1923; Hall, 1926; Faegri, 1937; Simpson,
Biologists might have been able to under- 1951, 1961). Others have treated them as
stand the philosophical issues and hence to units of a genetical nature (e.g., Huxley,
solve the species problem for themselves. 1940; Mayr, 1940, 1957, 1963; Camp and
However, the transfer of concepts across Gilly, 1943; Dobzhansky, 1950; Carson,
disciplinary boundaries from philosophy to 1957). Originally Dobzhansky ( 1935)
biology proved shallow to say the least. In treated them as classes of individuals, while
fact, the most elementary of philosophical Mayr treated them as classes of populations,
notions became garbled. "Organism" was but nonetheless the notion of a larger unit
confounded with "particular" because, at was there. Such genetical population con-
least in part, "individual" had designated cepts tended to grade into notions of species
both. However, more may be involved than as classes of genetically similar individuals.
just an equivocation. When someone uses A more extreme variant of the biological
a notion borrowed from an unfamiliar field species concept might be called the "organ-
of experience, he tends to confuse it with icist species concept." By this would be
a more familiar one which somewhat re- meant the notion that species are "super-
sembles it. An additional class of examples organisms." It asserts that species have
is available in what is sometimes called properties such as adaptation which do
"psychologism in logicm--confusing logical exist in organisms, but which have been
concepts with psychological namesakes or attributed to the higher level systems mainly
analogues ( Ghiselin, 1966a). by false analogy. Organicism is an old
The so-called "nominalistic species con- notion, one closely associated with a vitalist
cept" is based on the idea that classes are doctrine called "Holism" (for its history
not real, and if species are classes, it would see Ghiselin, 1974:29-30), which treats the
follow that species are not real. One way generality of classes as if they were com-
of answering it is to deny the basic premise posite wholes: as "aggregates" or individ-
and affirm that classes are real. However, uals. This view carries strong teleological
this is quite unnecessary, for species are implications, for if species are presupposed
individuals, and nominalists believe that to be cooperative units like organisms, then
individuals are real. The nominalistic spe- their component organisms would not com-
cies concept resulted from a garbled version pete, but would do things "for the good of
of thirteenth century philosophy. It is im- the species." Holism strongly influenced
pressive to see how many biologists have Emerson (1938) and other systematists and
supported it. To give a few examples: ecologists of the Chicago school, and is
Bessey, 1908; Britton, 1908; Lotsy, 1925; particularly evident in the later writings
Burma, 1949, 1954; Mason, 1950; Thomp- of Du Rietz (1930). It is largely responsible
son, 1952; Davidson, 1954; Cowan, 1962. for the group selectionist notions of many
Rebuttals have taken various forms. One evolutionists (e.g., Darlington, 1940). In-
way of answering it has been to say that deed, one can find a strong trace of it in
species are real in the sense that they are the writings of those who certainly would
SPECIES PROBLEM

not endorse the organicist species concept ristical value is that we find it much easier
were it formulated in explicit terms. to think of firms as individuals. For this
One good line of evidence suggesting that very reason a host of biological problems
Holism has been influential is the degree are more readily soluble if one learns to
to which intraspecific competition has been think like an economist. In addition we
deemphasized in favor of supposed advan- should note that we often fail to solve our
tages (often, but not always, to the species) problems because we cannot even identify
of maintaining populational integrity and them. Under such circumstances, concep-
specific distinctness. In his well-known tual investigations do more than just help.
book on species Mayr (1963) strongly de- They are the only way out.
emphasized sexual selection in favor of
isolating mechanisms in explaining sexual ACKNOWLEDGMENTS
dimorphism. Lately (Mayr, 1972) he has For comments on drafts of the manuscript, I am
changed the emphasis to a considerable grateful to Theodosius Dobzhansky, Patrick M.
extent, but some of us would feel that he Gaffney, David L. Hull, Ernst Mayr, David Wake
has not gone far enough (Ghiselin, 1974). and Leigh Van Valen. Support was provided by
N.S.F. Grant GS 33491.
Likewise, we find Mayr (1969b:316) saying
"The division of the total genetic variability REFERENCES
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