ARE BIOLOGICALSPECIES REAL?

*
HUGH LEHMAN
Iowa State University of Science & Technology

Difficultieswith the typologicalconcept of species led biologists to reject the "typo-

logical"presuppositionof an archetypewhich is manifest in each memberof a species.
The resultingconcept of species,which is here called the phenotypic species concept,
is considered as implying that biological species are not real. Modem population
thinking has given rise to the concept of a species as a gene-pool. This modem con-
cept is contrastedhere with the phenotypicconcept in light of some general criteria
for evaluatingspeciesconceptsand is shown to be more satisfactory.Finally, it is held
that to ask if a species is real is to ask whether the species grouping arrived at by
applying the principles involved in the species concept correspondswith groups of
organisms amongst which important biological relationshipsexist. It is argued that
in this sense species, as defined by the gene-pool concept, are certainly real.

1. Introduction. Contemporary biologists, such as Mayr ([7], p. 5) and Simpson

([8], p. 47) and others have stressed the importance for understanding evolutionary
theory of the rejection of "typological thinking" and "typological principles." The
basic concept of typological thinking is that groups of living organisms have (or
manifest) patterns which are common within any group to every member of the
group. The pattern has been referred to as the archetype or essence. The pattern
consists of a set of characteristics of organisms. Each member of the species shares
each of the characteristics of the essence of that species. Other characteristics of the
members of a species are considered, by the typologist, to be mere "accidents," that
is, unimportant. Typologists suppose that there is a hierarchy of archetypes having
the species as the lowest level. Mayr calls the concept of a biological species which
this type of thinking involves a typological concept of species. Sinlce the character-
istics supposedly shared in common by all members of the species were usually
morphological characteristics, he also calls this concept the morphological species
concept ([7], p. 16).
If one accepts the religious hypothesis that the species of living things were all
created by a special act of creation such that all organismsbelong to some species
and such that there are absolute discontinuitiesbetween distinct species, then one
can theoretically accommodate a typological species concept. However, given the
job of the practicing taxonomist and if one accepts the idea that the species evolve, a
typological species concept leads to some obvious difficulties.
Let us see how the idea that species evolve leads to difficulties. Perhaps all that
is required according to evolutionary theory in order for species to evolve is that
the organisms of a species are gradually modified in their characteristics so that
eventually the descendants of a given species are so different from their ancestors
that they belong to a different species from their ancestors. But, how could such
evolution occur? Given the typological concept of species, the accidental features

* ReceivedJanuary1967.

157

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158 HUGH LEHMAN

of organisms are unimportant. This would seem to imply that changes in these
features need not be taken account of in understandingevolution. An accumula-
tion of gradual changes in these features even leading to major modificationswould
be irrelevantto evolution. On the other hand, theoreticallyeven a small change in
an essential feature would constitute a new species. Thus evolution of a new
species out of an earlier species need not consist of an accumulation of gradual
changes; one small change would be sufficient. But this is clearly an unsatisfactory
notion of evolution if one is attempting to apply the concept of evolution to the
gradual emergence of new species of organismson the earth. Consider,for example,
the evolution of man. There is general agreement (see [31) that Homo Sapiens
evolved from the species Homo Erectus. But this evolutionary development does
not consist in just one small modificationof Homo Erectus. The characteristicsof
individuals of the species Homo Erectus might be possessed by some individuals of
Homo Sapiens and also perhaps by some Chimpanzees. There was a range of mor-
phological diversity amongst the individuals of Homo Erectus just as there is a
range of morphologicaldiversity amongst the individuals of Homo Sapiens. One
cannot find a distinctive Homo Erectus type which is shared by all and only Homo
Erectus individuals. It is possible that some individualsof the species Homo Erectus
differed as much from each other in regard to their morphologyas they would each
differ from an individual of the species Homo Sapiens.
A natural suggestion at this point is that the distinction between the essential
and accidental characteristicsof organisms be rejected. This might help to solve
the problems of the evolution of species. Further, this suggestion would seem to
be supported by consideration of taxonomical problems posed by the typological
concept. Taxonomy is concerned with the practical business of arriving at a syste-
matic classificationof organisms. Species, as we have indicated, would be the low-
est level of a hierarchyof typologically conceived categories. Species would belong
to genera etc. If the taxonomistis to arrive at a system of classificationthat can
be applied to the organismsthat exist (and which have existed and which will exist
in the future) he must base his categories on observed characteristicsof organisms.
But, in practice groupingorganismson the basis of their observed characteristicsdid
not lead to a satisfactorysystematic classificationin which the categories were de-
fined in terms of essences. In a great many cases one finds a continuous range of
organismswith no sharp discontinuitiesdividing the organismsinto discrete groups
on the basis of the characteristicspossessed by the individual organisms. Given this
situationit is naturalto take the position that any division of this range into seperate
species is arbitraryand further that there are often a number of different ways of
making such arbitrarydivisions. But taking this position is tantamountto rejecting
the idea that there is any difference in fact between the essential and merely acci-
dental characteristicsof an organism. Given one arbitraryclassificationsome ehar-
acteristics will be "essential"for some set of organisms while given another classi-
fication, equally arbitrary, other characteristics will be "essential" for those or-
ganisms.
Rejection of the distinction between the essential and accidental characteristics
of organisms has implied, for many people, that species are not real. Biological
reality was believed to consist of individual organisms and their parts. With re-
gard to most, if not all, of the characteristicsof organismsone could find a number

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 ARE RIOLOGICAL SPECES REAL? 159

of other organisms such that there was a continuous variation (within a certain
range) for any of these characteristics. A set of characteristicsmight be specified
in order to define a species but the basis of the specification was supposed to lie
entirely in the purposes of the biologist. Classification into species divided the
organismsinto distinct groups. But the species groupings did not necessarily cor-
respond to any "real' groupings of the organisms.
But what does it mean to say that species are not real, i.e., that species groupings
do not correspondto any real groupings of the organisms? Given the background
of the typological theory, real groups of organisms would consist in a number of
organismsthat were similar in all possessing the set of essential characteristicsand
which were absolutely distinct from any other such set, that is, in the case of
species, no other species would possess all of the characteristicsof the given species
and given any other species no organisms would exist between it and the origi-
nal species. To say that species are not real is to say that there are no such groups.
Given the theory that species gradually change into different species it is obvious
that during the course of time in which evolution of biological organismshas been
occurringthere are no such groups.
Given the acceptance of evolutionary theory, one may wonder why biologists
have not simply accepted the fact that species are not real as a fact of life. Of
course many biologists have accepted this as a fact. But to other biologists this has
not been an acceptable position. In many cases the species groupings arrived at by
biologists do seem to correspondto real groups of animals. If one considersa limited
geographicalarea within a limited period of time one can probably identify in that
spatio-temporalregion many groups in which the individual organismsare biologi-
cally related to each other in many different ways. They may be physiologically
and morphologically quite similar and also distinct in these respects from other
groups in the region. Members of such a group interbreed with each other. They
have common enemies. They may serve to sustain the existence of each other in a
variety of ways. Nearly any careful observer who entered such a region would dis-
cover the same groups. These facts seem incompatiblewith the theory that species
are not real. To many biologists the claim that species are not real seems to have
implied that there are no biological relationshipsthat unite organisms into definite
groups. Further, if all grouping is just arbitraryit seems surprisingthat any careful
observer would identify the same groups in a given spatio-temporalregion. Con-
sideration of the obvious reality of biological relationships between groups of or-
ganisms has kept open the hope that a satisfactoryconcept of species might yet be
arrived at, that is, a species concept which might reflect the grouping that do exist
in nature.
In the past thirty years biologists have defined a concept of species which is quite
different from the typological concept and which goes some way toward satisfying
the aim of those biologistswho have not been willing to accept the view that species
are merely arbitrarygroupings. In order to understandthis concept and to contrast
it with a typological concept whch has been divested of the distinction between
essential and accidental characteristicsit will be helpful to discuss some of the cri-
teria which are specificallyrelated by biologists to the definition of species.
2. Criteria for Criticizing the Concept of Species. In Biology the concept of a
species is a concept which may be used in the classificationof living organisms. (A

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160 HUGH LEHMAN

species however should not be considered as merely a classificatoryunit.) Species

units are included within larger units, namely genera, orders, families, kingdoms,
etc. and species have been divided into sub-units which have been called, varieties,
races, ecotypes, sub-species, etc. The different terms of sub-specific units do not
appear to reflect any systematic differences in the kind of group referred to, i.e.,
whether one calls a sub-specificunit a sub-species or a variety etc. is of no theoreti-
cal significance.
The term species is used in other areas besides biology. However, in many of
these other areas, calling a classificatoryunit a species, rather than a sub-species or
something else, is not significant. In these other areas, any classificatoryunit which
falls within a wider unit may be called a species unit. For example, a high school
student may be a species of student; and a Buick may be a species of automobile,
or a V-8 may be a species of automobile. For most purposes it would make no dif-
ference whether we called Buicks a species of automobile or a sub-species. Simi-
larly, it would make no difference for the classificationof high school students or
V-8's.
But the species concept of Biology is supposed to be applied in such a way that
calling something a species rather than a sub-species is significant. As we noted
above, the species category is believed by many to represent a clearly identifiable
level of biological organization. This category is more significant than either sub-
specific or super-specific categories. Ideally, the distinction between species and
sub-species on the one hand, and the distinctionbetween species and larger group-
ings on the other, should be fixed for all kinds of biological organisms. That is, the
way of making the distinction between species and sub-species (or species and
super-species) should not vary for different groups of organisms.
The importanceof fixing the meaning of the term species in biology can be seen
by considering certain statements that biologists have made. It has been claimed,
for example, that there are numerous species of some genera while in some other
genera there is only one or a few species. Thus, it is claimed that all human beings
belong to one species but that there are numerous species of paramecia. But how
are we to understandthe claim that there is only one species of man? It does not
imply that there are no distinguishablesub-groupsamongst human beings. Human
beings might be classifiedinto distinct sub-groupson the basis of blood types, facila
features, etc. Men, like paramecia, can be divided into distinct sub-groups. Why
are the sub-groupscalled species in one case but not in the other? For the above
statement about the numbers of species of human beings and paramecia to be un-
ambiguous, it is importantthat there be a concept of species which can be applied
to both human beings and to parameciasuch that the claim that human beings con-
stitute a single species is determined in light of the same criteria as determine the
claim that there are many species of paramecia. Let us call the criteria, which
determine whether a group of organismsis a species or not, species criteria; and
let us call the criteria which are used in criticizing species concepts species desid-
erata. In light of this last paragraphwe may then say that one desideratumfor a
species concept is that the criteria which determine species status are the same in
all cases. Let us call this the desideratumof species uniformity.
Not only must the same species criteriaapply to all biological organisms (so that
saying that paramecia aurelia is a species implies the same level of organization

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 ARE BIOLOGICAL SPECIES REAL? 161

amongst these organisms as is implied about human beings when human beings
are said to be a species) but these criteria must be such that all competetently
trained biologists who used them would arrive at the same classificationsof organ-
isms, if they had the same set of observationaldata to work on. For example, obser-
vations which would lead one biologist to say that some group of organismswas a
species should lead any other competent biologist to make the same judgment. This
is to say that species criteria should be objective. Preferences, tastes etc. of the
investigatorshould not enter into the factors which are relevant for deciding whether
a group of organismsis a species. Let us call this the desideratum of species ob-
jectivity."
Since Darwin it has also seemed important that species distinctions have some
evolutionary significance. Biologists are in general agreement that species distinc-
tions are irreversible. This is to say that if two groups of organismsbelong to dis-
tinct species then the successorsof these groups cannot belong to the same species.
Further the species is considered to be the smallest such irreversiblegrouping. If,
within a biological grouping, there are distinct sub-groupswhich cannot have suc-
cessors in common, then this biological grouping must be of higher order than spe-
cies. It must be sub-genera, or genera, etc. Thus, it would appear that a third
desideratumfor a species concept is that the species unit be the smallest irreversible
grouping of organisms. Let us call this desideratumthe desideratumof evolutionary
irreversibility.
Some people may feel that evolutionaryirreversibilityshould not be a desideratum
which is used to evaluate a species concept. Such people may feel that the evolu-
tionary irreversibilityof species is an empirical fact. The may think that this fact
has been discovered by the observationof existing or extinct species. However, this
does not appear to reflect the views of modern biologists. Ernst Mayr, for example,
says, "The primarycriterionspecies rank of a natural population is reproductiveiso-
lation ([7], p. 31). This criterion would guarantee irreversibilityof species.
The suggestion that the evolutionaryirreversibilityof species may be an empiri-
cal fact may be a result of thinking that the ideal division of organismsinto species
correspondsto some existing natural groupings. Natural, as it is used here, is op-
posed to artificial. Natural groupings were contrasted with groupings determined
with reference to human convenience. In practice, the search for natural groups
was a search for a way of classifying which did not separate inito different species
organisms in which certain key physiological functions were performed in nearly
identical ways. In recent years the idea of a naturalbasis of classificationhas largely
been given up. Where two organismsperform a given function in nearly identical
ways, this does not lead to their being put into the same species. The idea of con-
vergent or parallel lines of development is importantin explaining certain instances
in which different species of organisms are quite similar physiologically.2

1 In spite of the problem of attempting to achieve a concept of species which is uniform

and objective,the main business of classifying organismshas progressedwithout a great deal
of disagreement.Differentbiologists (and even non-biologists) who study particulargeographi-
cal regions usually agree on what species are present in such regions.
2 We should note that rejectionof the idea of natural groupsas species is not equivalent to
rejectingthe idea that species distinctionsare objective.

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162 HUGH LEHEMAN

A fourth desideratumfor a biological species concept is that species not be de-

fined with reference to specific spatial or temporal location. To take spatial or tem-
poral location as an essential feature of a given species would lead to the result
that organismsin differentlocations which were quite similar and which could have
descendents in common with the given species would nonetheless have to be classi-
fied as distinct species. There is, of course, a close connection between spatial loca-
tion and species distinctions. Spatial isolation is usually, if not always, the initial
causal factor leading to the division of a species into new species. Further, some-
times geographicalfactors enter into t:henaming of species. Let us call this fourth
desideratumthe desideratumof spatial and temporal irrelevance.3
3. The Phenotypic Concept of Species. In studying the work of biologists con-
cerned with taxonomyone finds several differentideas underlying classificationsinto
species. To explain these ideas several technical terms from biology may be help-
ful. At this point then, let us explain the terms genotype and phenotype.
It is generally accepted that the characteristicswhich any organism manifests
are a result of two factors. The first of these is the genetic structure which the
organism inherits from its parent or parents. The second of these is the environ-
mental influence which affects the developing organisms. The set of properties of
the developing and developed organism is called the phenotype of that organism.
The genetic structure which the organism acquired from its predecessors is the
genotype of that organism. The genetic structure of a given organism at a given
time is not necessarily the same as that of its predecessors or successors. A given
phenotype may be the result of several genotypes. In fact an importantproblem in
recent times has been to try to find out why the phenotypes of many groups of
organisms are as constant as they are given the number of variations in genetic
structurewhich occur within these groups.
It appears that we may formulate the phenotypic concept of species as follows:
A group of organismsis a species if and only if the members of the group are suf-
ficiently similar in phenotype. The phenotypic species concept is what results from
the typological concept once the assumptionthat each species representsan essence
or archetypeis rejected, i.e., once you give up the idea that there is any theoretically
significant distinction to be drawn between essence and accident.
Now let us critically discuss the phenotypic species concept. Suppose that we
have devised a way of classifying the organismsin a specific geographicalregion on
the basis of similaritieswithin groups as to phenotype and discontinuitiesbetween
groups. For example, suppose that all organisms in that region possessing charac-
teristics A, B, and C are lumped into one species while organisms lacking all of
these characteristicsare put into different species. If we could limit ourselves to
considering fairly limited and well-defined geographical regions, the phenotypic
species concept as illustratedhere would prove satisfactory. However, geographical
regions shade off into other regions, e.g., a valley between two mountain ranges
graduallyshifts from valley floor to foothills then to lower mountains and the valley

3 In light of the fact that thereare groupsof organismswhich are normallyseparated but
whichdo have commonoffspringwhen they comein contact,the fourthdesideratum is not
independentof the desideratum of evolutionaryirreversibility. on the
Speciesdistinguished
basisof spatialseparation couldhave commondescendants.

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 AIM BIOLOGICAL SPECES EAL? 163
may be connected to other valleys by mountain passes. Species ABC may exist as
a distinct species on the valley floor. But as we move further out from this point
we may find organismswhich fall into the discontinuitiesbetween the group ABC
and the other groups on the valley floor. The problem thus arises as to how these
intermediateorganismsare to be classified.
This example can illustrate several difficulties with the phenotypic species con-
cept. Since the species ABC is defined solely with reference to similarity or or-
ganisms in three respects there is no non-arbitraryway of handling the problem of
the intermediateorganisms. There would appear to be no more reason for dividing
ABC up into two sub-species, say ABC and AB not-C, than for calling the distinct
group a different species. To get on with the practical business of classifying or-
ganisms some arbitrarystipulationwould have to be made.
However, the fact that organismswhich do not completely fit into any one species
must be handled in an arbitraryfashion is not the main difficultyof the phenotypic
species concept. The main problem arises because the situation described in refer-
ence to group ABC is a problem which occurs frequently with many biological
groups. Thus arbitrarydecisions have frequently to be made for the purposes of
classificationand there is nothing in the phenotypic concept which provides a way
of making these decisions in a uniform manner in different cases.
No doubt, part of this problem in connection with the phenotypic concept of
species stems from the use of the vague expression"sufficientlysimilar"in the defi-
nition of this concept. How similaris sufficientlysimilar? One suggestion for deter-
mining the degree of similarityis to count the number of differences between or-
ganisms. But if this suggestion were to lead to the achievement of a uniform species
concept there would have to be some uniformly applicable way of deciding what
was a difference between organisms-a way which was uniform and applied to
plants and animals, micro-organismsand large organisms etc. While some success
has been achieved along these lines with groups of similarorganisms,no concept has
been developed which is theoretically applicable to all organisms.
Another suggestion for determining degree of similarity would be to determine
what sorts of traits of phenotypes were relevant as a basis for comparison. Species
might be defined with reference to some definite number of these relevant traits.
The relevant traits would be determined with reference to the importantbiological
functions such as reproduction. Historically,it appears that something like this has
happened in the classificationof species, for example, certain organismshave been
classified into species on the basis of similarities and differences in structure and
physiology of the reproductivesystem. However, the problem of the lack of a uni-
form species concept throughoutliving nature has not been solved in this way. The
features which seem relevant for classifying some groups do not seem relevant for
classifying others. Thus different features may be used depending on whether the
organism is a plant or an animal, reproduces sexually or asexually, is a micro-
organism,etc. Further, this suggestion does not provide the guarantee that species
groups will be irreversiblydistinct.
4. The Species as the Gene Pool. In recent times a definitionof species has been
suggested which is theoretically applicable to a great many groups of organismsif
not to all organisms. To explain this concept we must discuss the meaning of the
term gene pool. The idea of a gene pool is based on the idea of a breeding popula-

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164 HUGH LEHMAN

tion. A breeding population is a group of organismscontaining organismsof oppo-

site sexes where the probability of any pair of oppositely sexed individuals having
common offspringis the same (or nearly the same) as the probability of any other
pair. Since geographicalproximityobviously affects the probabilityof interbreeding,
breeding populations usually occupy one geographical region. But populations in
one region may occasionally interbreed with populations in other regions and the
offspringmay survive and join one or the other of the original populations. A gene
pool is defined as the most inclusive group of breeding populations between which
interbreeding sometimes occurs in their natural setting. (This is one definition of
gene pool. There does not appear to be complete uniformity amongst biologists as
to the use of this and related terms.) The concept of species we are now consider-
ing identified a species as a gene pool, thus we shall call this the gene pool concept.4
To help clarify the gene-pool concept we might first contrast it with a third con-
cept of species distinguishedby ProfessorMayr. Professor Mayr calls this concept
the non-dimensionalspecies concept ([7], p. 17). Accordingto the non-dimensional
species concept, two populations are distinct species if they fail to interbreed in
nature. (The qualificationin natureis importantin both these definitions. In nature
there are mechanismswhich prevent populationsfrom interbreedingwith each other
which are nullified to some extent in a laboratorysetting. There are cases of organ-
isms from distinct groups producing common offspring in the laboratorywhile in a
natural environment common offspring are not produced. Such mechanisms in-
clude factors which prevent mating from occurringand factors which prevent mat-
ing from being successful where it does occur.) But, two populationswhich do not
interbreed with each other may interbreed with still other populations which do
interbreed. Thus, if the non-dimensionalspecies concept is adopted by itself, it is
entirely possible in many cases for members of distinct species to have common off-
spring. Gene pools guarantee that the desideratum of evolutionary irreversibility
will be satisfied because there is no interchange of genetic material between dis-
tinct gene pools.
With regard to the desideratumof species uniformityit has to be noted that the
gene pool concept cannot be applied to all existing organisms. It can only be ap-
plied to populations of interbreedingindividuals. With regard to such individuals
it is much more successfulin relation to this desideratumthan is the phenotypic con-
cept of species. With regardto the gene pool concept, the term species conveys the
same biological informationfor a large number of different types of organisms. In
regard to the phenotypic concept, as we have seen, the factual content of the term
species varies in non-systematicfashion. With regard to the desideratum of objec-
tivity the gene-pool concept has a clear advantage over the phenotypic concept. It
has very often been noted that the decision as to whether a certain group of
organismsconstitutes a species is a matter of art when the phenotypic species con-
cept is involved. There are numerous cases in which a group called a species by
some biologists has been consideredto be some category other than species by other
biologists. The decision over whether a group of organismsis a species, using the
gene-pool concept of species, is not a matter of individual judgment.

4 ProfessorMayr calls this the multi-dimensionalconcept ([7], p. 19) This concept is multi-
dimensional because breeding populations are "distributedin the dimensions of space and
time."

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 ARE BIOLOGICAL SPECIES REAL? 165

In regard to the desideratum of evolutionary irreversibilityit is clear that the

gene-pool concept is more satisfactorythan the phenotypic concept. While dissimi-
larity of phenotype usually goes along with inability to interbreed,there is no guar-
antee that this will be so. Occasionallyclassificationsbased on phenotypic similarity
have been changed in light of informationconcerning interbreeding. But, two dis-
tinct gene-pools (species) can never contributeto the formationof a new gene-pool.
There are some difficulties with the gene-pool concept which should be men-
tioned. One of course, is that if species are gene-pools then some other classificatory
unit is needed for asexual or uni-parentalspecies. V. Grant has suggested that the
term binom be used for asexual organisms [4]. Since the term species is the tradi-
tional term of classificationthis suggestion has run into resistance from biologists
concerned with such organisms. Another suggestion is to retain the term species
with two meanings, one for asexual species and the other for sexual species. Pro-
fessor Mayr has favored this suggestion.
Grantingthe theoreticalimportanceof the gene-pool concept it may still be asked
why gene-pools should be identified with species. Why not retain the traditional
phenotypic species as the definition of species?
It appears that the reason for calling gene-pools species is two-fold. On the one
hand there is the dissatisfactionwith the phenotypic concept of species. This dis-
satisfaction stems primarily from the failure in regard to the desideratum of uni-
formity and also because species, as understoodphenotypically,do not relate to any
comprehensivebiological theory such as genetic theory. On the other hand, actual
classificationsof organismswill not be changed very much with the shift in species
concepts. Groups which were designated species in accord with phenotypic con-
cept will on the whole remain species. This latter point suggests that any attempt
to define the species concept might be tested to determine whether species con-
structed in accord with the definition in question actually coincide with the non-
theoretical classificationsdeveloped by naturalistsin the field.
The fact that the gene-pool concept allows for species which are temporallymulti-
dimensional would appear to create a serious difficulty for this concept. Breeding
populationsmay form spatial series. In a given region, population A may not inter-
breed with population B. But there may be gene interchange between population
B and population C in an adjacent region. And population C may interbreed with
Population D; and population D may in turn interbreed with population A. Ac-
cording to the gene-pool concept, these four populations all belong to the same
species. However, suppose we have a chain of non-contemporaneouspopulations.
That is suppose we have a chain of populations D, E, F, G,. . . Suppose that the
.

life-spans of the members of D and of E overlap and suppose that the life-spans
of the members of E and of F overlap and suppose that F and G similarly overlap
but that the lifespans of the members of D do not overlap with those of the mem-
bers of G. PopulationsD and G would belong to the same species since they belong
to the same gene-pool. But this leads to some surprisingand perhaps unsatisfactory
results. We can imagine a gene-pool with a temporal extent of hundreds of thou-
sands or even of millions of years. But the populations at widely separated places
on these temporalchains might be as widely disparateas eohippi and modem horses
or as gorillas and modem men. Clearly, to avoid such absurd consequences, the
gene-pool concept of species needs to be modified in some way.

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166 HUGH LEHMAN

One suggestion for modifying the gene-pool concept is to require that all the
populationsin a gene-pool which make up a species be contemporaneouswith each
othier,that is, each population must temporally overlap with all other populations
in the gene-pool for the group of populations to constitute a species. This sugges-
tion does not seem very satisfactory. It would violate the desideratumof temporal
irrelevance. Thus, for example it would lead to the consequence that the human
beings who fought the American Revolution would belong to a different species
than that of human beings living today. We would be separated not only in cul-
ture from the Atheniansbut in species also.
This problem has been recognized by biologists. As we have indicated a multi-
dimensional species would lump temporally continuous chains of populations into
one species. One could start with modem man and work back through all the popu-
lations in the chain. The obvious solution to this problem is to attempt to develop
a uniform phenotypic species concept which can be applied so as to divide such
continuous populations into distinct species. Along these lines, Simpson has sug-
gested that "Successivespecies should be so defined as to make the morphological
difference between them at least as great as sequential differences among contem-
poraneousspecies of the same group or closely allied groups." ([81, p. 165) Thus,
Homo Sapiens would be distinguished from other groups in the temporal chain.
We would belong to the same species as the Athenians because as far as we know
the phenotypes of the Athenians would fall within the limits of the variations that
exist among Homo sapiens at the present time. We would be distinguished from
Homnoerectus in species because the phenotypes of these individuals do not fall
within the range of contemporaryvariations of phenotype. The necessity to take
this step clearly leads to assume sacrifice of objectivity.
5. Conclusion. In conclusion let us briefly refer our discussion of the gene-pool
concept of species back to our original question concerning the reality of species.
To ask if a species is real is to ask whether the species grouping arrived at by apply-
ing the principlesinvolved in the species concept correspondswith groups of organ-
isms amongst which importantbiological relationshipsexist. The species, as defined
by the gene-pool concept, is certainly real in this sense. Membershipin a common
gene-pool unites organisms into a group and separates them from other groups.
Certainly, phenotypic species concepts divide groups of organisms into units also.
However, the gene-pool evolves as a unit. The genotypes of one gene-pool cannot
be contributedto the genetic material of another gene-pool.5 What happens to the
members of a gene pool affects the genetic potentialities for future generations of
the same gene-pool but does not, in itself, affect these potentialities for other gene-
pools. (Of course, what happens in one gene-pool may affect other gene-pools also
since the survival or non-survivalof the members of one may affect the survival or
non-survivalof members of the other.) Species defined without regard for gene-
pool do not necessarily evolve as a unit. For example, the death of some members
of such a species may not directly affect the genetic material contributed to subse-
quent generationsof other members of the species. This would happen if a species
included more than one gene-pool. Or consider that a mutation which occurs to a
Chimpanzee will be contributed to the gene-pool from which other Chimpanzees

5 See Dobzhansky, [3], to whom this argumentis largely due.

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 ATIE SIOLOCICAL SPECXIS REAL? 167

are formed but will not be available for the formation of any human beings. A
phenotypic concept of species might lead to a division of organisms into two dis-
tinct species which organismswere members of one gene-pool. In this case each
of the two species would not evolve as a unit. Genetic material from one species
could be contributedto the formation of new members of the other species. Such
species would be like races.
We saw, in our discussion of the typological concept, that a typological species
concept led to problemswhen one attempts to understandevolution. To accommo-
date the facts of evolution in an adequate theory which enables us to understand
evolutionaryphenomena, the reality of species had to be abandoned. However, the
gene-pool species concept does not lead to these problems. A species, conceived
as a gene-pool, can undergo change in various ways. The distributionof genotypes
at a given time can change. Over a sufficientlylong period of time the phenotypes
and genotypes associated with a multi-dimensionalspecies can undergo profound
modifications. Since a gene-pool species can undergo change while continuing to
be the same species, the gene-pool concept of species has been called a dynamic
concept ratherthan a static concept. The modificationsof species are neither incom-
patible with the unity of a gene-pool species nor with its distinctness from other
species. Thus, the gene-pool concept of species does not produce theoretical prob-
lems such as those which arose from the attempt to understand the evolution of
species when species were understood typologically.

REFERENCES

[1] Dobzhansky,Theodosius, "A Critique of the Species Concept in Biology," Philosophy of

Science,Vol. 2, No. 3, July, 1935, pp. 344-355.
[2] Dobzhansky,Theodosius,Evolution Genetics and Man, (New York:John Wiley and Sons,
Inc., 1955).
[3] Dobzhansky,Theodosius, "Genetic Entities in Hominid Evolution" in Classificationand
Human Evolution.
[4] Grant, V., "The Plant Species in Theory and Practice,"in [6], pp. 39-80.
[5] Mayr, Ernst, Biological Systematics,Sixteenth Annual Biology Colloquim, April 30, 1955,
Oregon State College, Corvallis, 1955.
[6] Mayr, Emst, Ed., The Species Problem, A SymposiumPresented at the Atlanta Meeting
of the AmericanAssociationfor the Advancementof Science, Publication No. 50, 1957.
[7] Mayr, Ernst, Animal Species and Evolution, (Cambridge,Massachusetts,HarvardUniver-
sity Press, 1963).
[8] Simpson, G. G., Principles of Animal Taxonomy (New York: Columbia University Press,
1961).

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All use subject to University of Chicago Press Terms and Conditions (http://www.journals.uchicago.edu/t-and-c).