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Competition theory, evolution, and the concept

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Article in Acta Biotheoretica February 1982

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 Acta Biotheoretica 31,165-179 (1982) 0001-5342/82/030165-15 $00.20/0
1982 Martillus NijJIO!!/Dr W. Junk Publishers, The Hague. Printed in the Netherlands

Competition theory, evolution, and the concept of

an ecological niche

THOMAS R. ALLEY

Growth Center, L-7063, School of Dental Medicine, University of Connecticut Health

Center, Farmington, CT 06031, USA

(Received I5-VIl-198})

Abstract. This article examines some of the main tenets of competition theory in light
of the theory of evolution and the concept of an ecological niche. The principle of
competitive exclusion and the related assumption that communities exist at competitive
equilibrium - fundamental parts of many compctition theories and models - may be
violated if non-cquilibrium conditions exist in natural communities or are incorporated
into competition models. Furthermore, these two basic tenets of competition theory
are not compatible with the theory of evolution. Variation in ecologically significant
environmental factors and non-equilibrium in population numbers should occur in most
natural communities, and such changes have important effects on community relations,
niche overlap, and the evolution of ecosystems. Ecologists should view compctition as
a process occurring within a complex dYllamic system, and should be wary of theoretical
positions built upon simple laboratory experiments or simplistic mathematical models.
In considering the relationship between niche overlap and competition, niche overlap
should not be taken as a sufficient condition for competition; many factors may prevent
or diminish competition between populations with similar resource utilization patterns.
The typically opposing forces of intraspecific and interspecific competition need to be
simultaneouslY considered, for it is the balance between them that 1n large part
detcrmines niche boundaries.

1. Introduction
The concepts of an <ecological niche' and of 'competition' are of great
importance in modern ecology and population biology. Moreover, niche
theory and competition theory have been closely associated since the concept
of an ecological niche first appeared near the beginning of this century. Yet,
as this paper points out, these two are often used in connection with each
other in a manner that entails incompatibility with each other, with observa-
tions of natural communities, or with one or more fundamental tenets of
evolutionary theory. In addition to gathering together such criticisms of com-
petition and niche theory, this article attempts to suggest ways to avoid them.
Before turning to these tasks, it is necessary to briefly defme both niche and
competition.
Deji""nitiolls of 'ec%gica/niclze' and 'competition'
The tenn 'ecological niche' or 'econiche' has been used to refer to a variety of
concepts (see Hutchinson, 1978; Vandermeer, 1972; Whittaker and Levin,
1975). It will be used herein as a functional concept; an ecological niche
consists of the conditions necessary to support the vital activities of a type of
organism. An ecological niche can be thought of as the dual complement of

166
an organismic unit (such as a species or individual) when an organismic unit
is defined by some degree of functional equivalence; that is, an ecological
niche is the dual complement of a jil1lctionally specified la,:'(onomic unit
(FSTU) (Alley, 1982). The niche of a FSTU is defined by the range of
environmental variation in both biotic and abiotic factors under which that
organismic unit can engage in the activities necessary for its survival. Thus a
niche can be viewed as a 1l1!lltidimensional system of relations between a
FSTU and the environment (Rejmanek and Jenik, 1975), and may be repre-
sented by a volume in a multidimensional niche space within which an
organismic unit's responses are inside the critical boundaries for survival (see
Wangersky, 1972; Wuenscher, 1969).
The niche concept can only remain clear and retain much of its usefulness
(such as for comparison of similar roles in different communities) if some
'blurred vision' is employed so that 'similar' functional relations can be identi-
fied (MacArthur, 1968). The niche concept need not, and should not be
strictly tied to species as the unit of analysis. It is often useful to treat two or
more organismic units as functionally equivalent (Le., as having the same niche
or 'profession') even though their niches are not absolutely identical. Ecolo-
gists typically ignore relatively minor functional differences between organ~
isms in referring to a niche of an aggregate of organisms, or a niche of one
or more organisms over" an extended period of time (Alley, 1982 ~ MacMahon
et ai., 1981). In this manner, equivalency can be seen across organismic units
below, at or above the species level, and captured in an appropriate niche
specification, even though their niches will not prove to be identical should
enough niche dimensions be examined in sufficient detail. This may be done
regardless of phylogenetic relationships, as is done in conceiving of 'eco-
species' (or 'ecological equivalents') (see Alley, 1982; Odum, 1959). Elton's
goal of recognizing cross-species functional equivalency need not, as some
ecologists (e.g., Williamson, 1972) have claimed, be forfeited with acceptance
of the multidimensional niche.
In this paper, mutual exploitation of at least one limited resource will be
seen as a necessary (but not a sufficient) condition for competition. Competi-
tion is a resource-related form of interaction between two or more organisms
(Birch, 1957; Klomp, 1961). Competition should be understood in connection
with the niche concept since some similarity -or overlap in niches is a pre-
reqUisite of competition, and since niche shifts provide the clearest evidence
of competition (Diamond, 1978).
2. Competition theory and ecological niches
The views of many ecologists on the nature of niches are beset with a number
of difficulties related to competition. In large part these problems concern
two basic and Widely accepted propositions of competition theory: the prin~
ciple of competitive exclusion - which asserts that there can be no more
167
than one species per niche (or per limited resource) - and the corollary that
communities exist at competitive equilibrium (see Cole, 1960; Hardin, 1960;
Levin, 1970; Miller, 1967). Although current niche theory is, in large part,
closely tied to these propositions, they have become increasingly suspect
in recent years.
Competitil1e equilibrium
To begin, many theorists and 'virtually all mathematical treatments of com w
petition' assume competitive equilibrium (Wiens, 1977), but there is growing
evidence that true competitive equilibrium occurs rarely, if at all, in natural
situations (Huston, 1979; Pianka, 1976). This should come as no surprise
since, contrary to the prevalent view of earlier naturalists, it is clear that
organisms do fluctuate greatly in numbers, and not simply because of develop~
ments in human civilization that interfere with 'natural' conditions (den Boer,
1979~ Elton, ] 950; Odum, 1959; Wangersky. 1978). Indeed, it is appropriate
to refer to some species, such as the Paramecium living in a puddle that will
soon dry up, as 'non-equilibrium species' (Leigh, 1971), for these species
opportunistically exploit spatially discrete, temporary sets of environmental
conditions, but will eventually be subject to extinction in that locale. Cyclic
climatic and tidal variations of various periodicities, geological (e.g., volcanic
eruptions), climatic (e.g., late frosts) and organic (e.g., viral or bacterial
infestations) irregularities, migrations, invasions and other changes in the
distributional patterns of biota, fires, alterations in population gene pools,
and oscillations in the numbers, status (e.g., metamorphic state), and activities
of many organisms will all ensure that natural conununities do not maintain
states of competitive equilibrium: the only likely exception being communities
that form an extremely simple, homogenous, and relatively closed ecological
system, such as those found within geothermal irregularities. Levins (1979)
has more formally shown that a community with temporal variation and
nonlinear dynamics (as arise from learning and seasonally variable feeding
rates, among other things) may persist without reaching a stable equilibrium.
Even if the underlying tenets of the competitive exclusion principle were
true, equilibrium is not expected to occur in most organisms with generation
times of days or weeks, such as plankton and numerous insects (Hutchinson,
1961); nor is it to be expected when members of two sympa tric and compe-
titive species inhibit their own population growth more than they inhibit that
of the other species (Cole, 1960; Stewart and Levin, 1973). Finally, the niches
of some species seem to include niches of other species, with the included
species apparently favored in the more optimal (for both species) parts of
the joint fundamental niche space, and the species with the broader realized
niche favored by greater adaptability to environmental variation (Colwell and
Fuentes, 1975; Hutchinson, 1978; Miller, 1967). Under these circumstances,
interspecies competition is almost certainly a necessary consequence of niche
overlap. (The relationship between competition and niche overlap is discussed
further in the final/of this paper.)
s~ctjM
168 169

Competitive exclusion (Miller, 1967, p. 4). Simple experimental studies may elucidate ~he elementary
processes that govern community organization and. e~olu~lOn~ but :hey
Without competitive equilibrium, it is unlikely that strict competitive exclu-
cannot be expected to solve the mysteries of species dlstnbutlon In the field,
sion can be maintained. Although there are a large number of competition
niche differentiation, or organization and succession of natural communities
models for two populations, they all (except one: de Bruyn, 1980) support
(McIntosh, 1970).' Furthermore, while the laboratory experiments have
the principle of competitive exclusion by predicting eventual extinction for
typically concerned the interactions of only two species, and while such
one of the populations, given constancy in e11l l ironmental conditions and
simple ecological models as the Lotka-Volterra equations (also see Levins,
genetic composition. In accordance with these models and the principle of
1968; May, 1973) assume th at in teractions between any pair of species are
competitive exclusion it is commonly assumed that 'the coexistence of any
independent of the species comprising the rest of the community, competition
two species, no matter how similar they might seem, implies that the species
(and other factors, such as predation, that determine realized niche dimensio~s)
are not in true competition' (Wangersky, 1978, p. 203), but non-equilibrium
has been found to vary with the presence or absence of (an)other speCIes
conditions in natural environments or populations can produce violations of
this assumption. When temporal variations in environmental conditions are (Colwell and Fuentes, 1975; Diamond, 1975; MacArthur, 1972; Neill, 1974;
incorporated into competition models, coexistence of competitors becomes Pianka, 1976). Actual ecological niches are 'nor necessarily, and perhaps
a significant possibility (see Grenny, Bella and Curl, 1973; Koch, 1974). hardly ever, a simple interaction of pairwise species interactions' (Colwell
Stewart and Levin (1973) found that with seasonal variation in resource and Fuentes, 1975, p. 300).
exploitation it is possible to obtain 'stable, nontrivial equilibrium' from a The evidence for competitive exclusion in natural populations is at best
model for two species even if they are to exist on a single resource. Likewise, 'spotty and often almost anecdotal' (Wangersky, 1978), and it is likely .that
empirical studies indiate that even slight fluctuations in a single environmental complete competitive exclusion occurs rarely outside the laboratory (BlICh,
factor (e.g., temperature) can be sufficient to reverse a competitive advantage
1957, 1979; den Boer, 1980; Hutchinson, 1961; Miller, 1967; Roughgarden,
and may permit indefi~ite coexistence of competing species (den Boer, 1980;
1976; and below). As Gilbert et al. (1952) pointed out, the decline of a
population due to competition will always take some time, and if the rate of
Hutchinson, 1978; Kaplin and Yorke. 1977; Klomp, 1961; Miller, 1967;
decline is sufficiently slow, genetic change may spread through the popula-
Schoener, 1977). For example, when provided with adequate soil nutrients
tion so as to decrease or even reverse the course of competition. 'Species
the annual grassland species Bromus molUs appears to be competitively
are not static entities with fixed relations to the environment, but plastic
superior to its competitor, Erodium botrys, but the latter is favored in drought
elements, changing their genetic constitution under the influence of the
years (McGown and Williams, 1968). Some recent mathematical simulations
physical factors of the environment and of the interactions with other
(de Bruyn, 1980) indicate that coexistence of pairs of competitors is possible
species' (Lewontin, 1968, p. 3). Furthermore, when adverse conditions arise
even without environmental fluctuations if there are at least two density
due to competition, a behavioral change may appear that will alter the
limiting factors (such as a single shared resource and either intraspecific
competitive relationship between populations (Gilbert et al., 1952). ~lear
territoriality or density dependent predation) per pair of competitors. To the
experimental evidence of such a reduction in competition through behavlOral
extent to which the various types of coexisting organisms are limited by
plasticity has recently been provided by Werner and Hall (I976). They
shared resources, the fluctuations in the numbers of one type of organism can
examined the food habits of three naturally coexisting species of the sunfish
be expected to have a profound effect on others in the same community: an
genus Lepomis when stocked either separately or together. In all cases they
effect opposing the stabilizing processes which might otherwise lead to the
were stocked in replicate environments consisting of small circular ponds.
establishment of competitive equilibrium.
These species display quite similar resource utilization patterns when stocked
Although some laboratory experiments with simple, homogenous environ-
separately, but because of changes in the optimal patterns of resource utiliza-
ments have demonstrated complete competitive exclusion (see Hardin, 1960),
tion, each species engages in a distinct pattern of resource utilization when
these experiments should be seen in retrospect as instructive 'because of their
the other two congeneric species are present.
con trast with nature rather than their parallel to it' (MacArthur, 1972, p. 25,
Even if competitive exclusion does occur, reintroduction may often be
emphasis added); they demonstrate that some minimal environmental fluctua-
possible, for no ecotope is a completely closed biological system. Moreover,
tion or heterogeneity is sometimes required for coexistence. As a review of
species that have inhabited a locale in the past are more likely to undergo
the literature on competition concluded, 'in their anxiety to "prove" the
successful reintroduction than are species that have not lived under those
competitive exclusion principle, ecologists have repeatedly ignored the impor-
ecological conditions (Hutchinson, 1978), especially if, the former c.ontinues
tant fact that many, if not most, laboratory studies of competition have
to live in neighboring areas in which it has an ecological advantage (NIcholson,
provided better evidence of coexistence than ... of competitive exclusion'
1960). The large role played by chance in determining the seed or egg dis-
170
persal patterns in many species may prelude effective selection for mutual
exclusion by competition.
3. Competition and evolution
Although the competitive principle and the assumption of competitive
equilibrium arc helpful in simplifying models of competition and species
diversification, and seem to be applicable to some simple and homogeneous
environments, they are not consistent with evolutionary theory. Three main
points of inconsistency should be noted.
First the idea of a trend towards ever increasing equilibrium does not
mesh w'ith the dynamic concept of evolution, or with current theories of
factors governing extinction and phylogenetic divergence. It is generally
assumed that phenotypic evolution of communities goes on forever without
reaching a steady state (e.g., Lewontin, 1979; Wangersky, 1972). Indeed,
following Waddington (1969), it can be argued that the combined effect of
organismic modification of the environment and evolutionary modification of
organisms is an ever increasing rate of environmental change associated with
an increasing rate of change of selection pressures for evolutionary modifica-
tion: the environment which exerts selection on one living organism is influ~
enced by the pressures of other living systems, and as these other organisms
change in evolution, so too does the environment of the first organism, and
it must evolve as well, entailing further environmental changes and so on. In
tlus way, organic evolution engenders a fonn of positive feedback wherein
change itself generates change, resulting in an ever accelerating rate of change.
Extinction of a species is often (perhaps too often: see Birch, 1979;
den Boer, 1980; and above) attributed to the coexistence of other competi-
tive spec"ies; indeed, competition may be 'one of the commonest initiators of
extinction' (Simpson, 1967, p. 206), at least on islands (MacArthur, 1972).
While forms of phylogenetic divergence such as character displacement (Brown
and Wilson, 1956; Lack, 1971) may well reflect a tendency towards competi-
tive equilibrium, natural selection would not have produced such effects
unless competition had a significant and long-term effect on reproductive
success. Similarly, adaptive radiation is generally attributed to the selection
pressure from intraspecific competition (Lack, 1971). As argued above, .it
seems unreasonable to maintain that species can always manage to aVOId
competition or, as Lack (1971, p. 8) seems to suggest, that (interspecies)
competition has occurred in the past but no longer arises thanks to the
equilibrium provided by competitive exclusion. Both intraspecies and inter-
species competition have been clearly shown to affect a wide range of organ-
isms and to be major determinants of the organization and population densi-
ties of natural communities (Bakker, 1964; Colwell and Fuentes, 1975;
Connell, 1975; Cox et aI., 1976; Knight-Jones and Moyse, 1961; MacArthur,
1972; McNaughton and Wolf, 1970; McIntosh, 1970; Schoener, 1977; Stem
and Roche, 1974).
171
Second, given a view of evolution as the movement of a niche through
niche space such that reproductive success tends towards optimization (Le.,
to see any actual niche as the result of a never-ending evolutionary 'search'
for an optimal way of relating to an environment: see Lewontin, 1978;
Wangersky, 1972), it seems reasonable to expect that more than one species
might move towards the same niche space; i.e., that overlapping regions of
niche space could provide roughly optimal affordances for more than one
species. (This is not equivalent to a view of evolu tion as adaption to niches
which, as Lewontin (I 978) convincingly argues, leads to many difficulties
with the concepts of both adaptation and ecological niche.) Moreover, the
existence of both parallel and (especially) convergent evolution provide
evidence that the adaptive possibilities in a given type of environment are
rather narrowly limited (see Mooney, 1975). Independently evolving species
under similar environmental conditions have often responded to sin1ilar
selection pressures with nearly identical adaptations, sometimes becoming
similar enough to be called 'ecological equivalents' (see Cody, 1973b; Cox
et al., 1976; Pianka, 1974; Sage, 1973). Even within the same geographical
area, different species may display character convergence indicative of conver-
gence on niche dimensions. Such convergence is seen in tropical rain forest
trees of the same stratum which are often hardly distinguishable in leaf and
crown characters despite membership in different species, and even different
genera (Richards, 1969). Recent evidence indicates that even with notable
morphological differences, coexisting species may respond in a similar fashion
to resources in their environments (Wiens, 1977).
Third, and this is perhaps the Malthusian/Darwinian insight, replication in
a finite (and, therefore, resource limited) environment will eventually lead to
competition and natural selection (see Milne, 1962). In the words of M.T.
Ghiselin, 'the economy of nature is competitive throughout, and selection,
whether natural or sexual, is the most compelling manifestation of this truth.'
(1974, p. 57). Intraspecies competition has often been ignored in competition
models, but is a well-accepted aspect of natural and sexual selection. It also
seems possible that some organisms under certain circumstances can increase
their fitness by engaging in interspecific competitive behaviors, as in inter-
specific territorial behavior (Grians and Willson, 1964; Verner, 1977).
Schoener (1977) has noted some potential benefits for Reptilia engaging in
interference competition.
None of these shortcomings of either the principle ofcompetitive exclusion
or the notion of competitive equilibrium necessitate rejection of the view that
populations can and will increase their fitness by 'following' ontogenetic and
phylcrenetic 'paths' that limit competition. Nevertheless, interspecific com-
petition may not be a sufficient condition for movement towards avoidance
of competition. First, natural selection should comprise between pressures
for ecological divergence due to intraspecific competition and ecological
fixity due to disadvantages inherent in shifts to different resource utilization
172 173
Thus, the selection pressures stemming from interspecific competition
(Roughgarden, 1976). Klomp (1961) has suggested that most cases of stable
may often be insufficient to lead to competitive exclusion, for the width
polymorphism can be attributed to intraspecific competition. Second, as
and overlap of niches should tend toward equilibria between the selection
illustrated in Figure 1, intraspecific and interspecific competition can be
expected to have opposite effects with regard to resource utilization, with pressures produced by intraspecific and interspecific competition. The relative
intraspecific competition generally acting to broaden niches and interspecific pressure exerted by intraspecific versus interspecific competition may,
competition tending to make niches narrower (Svardson, 1949). however, vary as competitive relations change during a successful sequence
/- - - - '\. ... ' ..... (McIntosh, 1970): a process which would limit attained degree of competi-
tive equilibrium. In any case, competition is by no means the only deter-
/ .'000 .. _ minant of evolutionary selection (Birch, 1957; Darwin, 1859; Milne, 1961),
/ "-" =\ -0 0 and neither econiches nor population equilibria should be assumed to be solely
I ~:. ~ 0'0 a result of competition (Gilpin and Case, 1976~ Wiens, 1977).
A theory of limiting similarity (e.g., MacArthur and Levins, 1967;
t
~ "00. /
./ 0 /
: '
MacArthur, 1972) due to a tendency to avoid interspecific competition may
I
t
~ /
/' \'00. c.c :

be acceptable, but the theory of competitive equilibrium due to absolute
exclusion of competitors is not. Evidence of limits on niche similarity of
J ~ / '- ) .. '. ,/ =/ - " coexisting species can be found (e.g., Hutchinson, 1978; Lack, 1971; Pianka,
I ~/ '" /O'o~ I 1976; Schoener, 1974; 1977), but the actual niche, habitat, and range bound-

/ ~ /~ ~~ ~ \ aries of species will often overlap and should not be expected to be fixed (see
Emlen, 1973; Grant, 1981; McNaughton and Wolf, 1970; Wiens, 1977). Even
/ --- I I when ecological niche differentiation can be demonstrated, competition may
~\ J still arise due to resource limitations in the exclusive niche regions. For
example, even though species of European tit (Pan's) prefer different types
./' --J~ I of nesting sites, at least j'il'e species will compete for a single type of site

~1 . /1~) ~
(Lack, 1971). As a more striking example, in a number of phyla coexisting

0
0
,..:-
~--- /
/
; species of quite different sizes have been found to consume highly similar
types and sizes of prey (Wiens, 1977; Wilson, 1975). Complete ecological
isolation of organismic units is unlikely without habitat or range separation.
While it is quite reasonable to expect no two coexisting species (or even
smaller taxonomic units; see Alley, 1982; Macmahon et al., 1981) to have
t l' \_/
,
precisely the same niche structure, and to expect the niche overlap of eco-
logically similar or closely related species to be inversely related to overlap in

...... . . .. . ... o geographical are"a, the competition and disequilibrium which remain will have

.... ..-.... a profound influence on ecological relationships and on the evolution of

Figure 1. A schematic two-dimensional diagram depic'ting opposing forces on a species'
(or population's) niche boundaries. Intraspecific competition creates pressure to increase
niche breadth, while competition from five other species creates a countering pressure
for niche contraction. The resultant boundaries for a species are the balance between
intraspecific and interspecific competition.
As expected from this perspective, species' 'ranges of adaptability' (niche
breadth) seem to be positively correlated with the degree of interspecific
competition encountered (e.g., Bernstein, 1979; Mooney, 1975). The typically
great degree of niche similarity among conspecifics adds to the power of the
selective pressures for niche expansion produced by intraspecific competition
(McIntosh,1970).
ecosystems. Ecosystems may exhibit relatille equilibria, but it is crucial for
both ecological and evolutionary theory that this not be mistaken for or
treated as absolute equilibrium.
4. Niche overlap and competition
A common problem in current econiche theory is a tendency to equate niche
overlap and competition, such as in Odum's statement that 'competition
occurs whenever niches overlap even to a partial extent' (1959, p. 231). In
fact, as Pianka (1976) points out, measures of niche overlap have often been
used to estimate the competition coefficients in Lotka-Volterra style compe-
174
tition models (e.g., Cody, 1974; Levins, 1968; MacArthur, 1972; May and
MacArthur, 1972). Using measures of niche overlap and competition inter-
changably, however, implies a failure to recognize that niche overlap need not
entail competition. While it is well-recognized that niche overlap will not lead
to competition if potential competitors have discrete (allopatric) geographical
distributions, it is too infrequently realized that other factors can mitigate or
preclude competition. Indeed, competition and niche overlap may sometimes
be i/ll'ersely related (Pianka, 1974; 1976; Wiens, 1977).
Competition should not occur if the relevant resources in the region of
niche overlap are not in short supply; interspecies competition is not likely
to occur unless two (or more) sympatric populations actually share a popu-
lation-limiting resource. Climatic factors, parasites, predators, density-
independent resource shortages, and disease can (theoretically, at least)
reduce fecundity, fertility, and longevity, so effectively limiting population
numbers that competitors (or potential competitors) can coexist; there is
substantial evidence of the effectiveness of these and other density-independent
factors in limiting pop ulations (e .g., Connell, 1975; Geisel, 1974; Harrison,
1964; Hutchinson, 1978; Kendeigh, 1961;Miller, 1967;White, 1978). When a
population-limiting factor is especially effective on a competitively dominant
species. other competitively inferior species may be enabled to share its
niche space.
As Hutchinson (1957) carefully poin ted outin presenting his (set-theoretic)
hypervolumetric conception of the econiche, set-theoretic models fallaciously
presuppose that all points within a niche space are equally suitable for the
organism in question. Actual econiches, however, will contain an optimal part
surrounded by suboptimal parts nearer to 'cloudy' or fuzzy niche boundaries
(see Emlen, 1973; Giesel, 1974 ; Miller, 1967; Stern and Roche, 1974). Even
if the niches of two competing species (or populations) completely overlapped,
all that would be required for coexistence is that each be a superior competi-
tor in a different subregion of the niche hyperspace, whether by virtue of a
greater efficiency in exploiting resources, or by more effective direct (inter-
ference) competition (Colwell and Fuentes, 1975).
The distinction between relatively direct or 'interference competition'
(including territorial behavior) and indirect or 'exploitation competition',
such as that stemming from mutual use of a limited resource (Colwell and
Fuentes, 1975; Miller,. 1967; Pianka, 1976; Schoener, 1977), permits addi-
tional clarification of the relationship of niche overlap to competition. Not
surprisingly, genetic relatedness is associated with niche overlap (or ecological
similarity; see, e.g., den Boer, 1980), especially in the case of conspecific
individuals. For this reason, exploitation competition (and probably inter-
ference competition as well; Schoener, 1977) is much greater within than
between species. A less obvious point is that niche overlap 'is neither a
necessary nor a sufficient condition for interference competition; moreover,
overlap is only a necessary but not a sufficient condition for exploitation
175
competition' (Pianka, 1976, p. 122). Under some circumstances, a small
degree of overlap (e.g., mutual use of a limited food source) can result in
exploitation competition, but even food shortages do not necessarily entail
competition (White, 1978). Finally, overlap in the niches of mature members
of different species may be associated with little or no interspecific competition
if no overlap is present in earlier development stages (Stern and Roche, 1974).
It can be expected that evolutionary selection will generally tend to
minimize exploitation competition (Christiansen and Fenchel, 1977; Gilbert
et al., 1952). The consequences of competition may include microhabit
divergence (as in some pairs of flatwonn (Planaria) species; Beauchamp and
Ullyatt, 1932), or habitat shift (as in many related bird (Lack, 1971) and
reptile (Schoener, 1977) species), competitive exclusion producing differen-
tia tion of geographic distribution (as in the Lactera lizard species studied by
Nevo et al., 1972), adaptive radiation (as in Darwin's finches: Lack, 1971;
Grant, 1981), range restriction (as in several coexisting species of shorebirds
that have similar diets but forage at different distances from shore; Cody,
1973a), ecological variation via polymorphism (as in many cases of sexual
dimorphism; Selander, 1966), or the displacement (as in character displace-
ment) or compression (see Christiansen and Fenchel, 1977; MacArthur, 1972)
of econiches. Furthennore, increasing competition (either intraspecific or
interspecific), insofar as it is density dependent, will tend to shift life-history
strategies towards K selection (a strategy of producing offspring of high
competitive ability [fitness] on the r-K continuum; in the (idealized) case
of completely density independent population control in which there is no
competition, r~selection (i.e., maximization of r, the intrinsic rate of natural
increase) will be favored (MacArthur and Wilson, 1967; McNaughton, 1975;
Pianka, 1970). These effects are not mutually exclusive, and may be more
directly a result of (potential) interference competion than of exploitation
competition.
5. Conclusion
The foibles and fallacies in the literature on competition and the niche
contain a moral for niche theory: a viable defmition of the niche needs to
be based more on ecological data from natural communities, and less on
idealized ecological models and data from highly artificial laboratory experi-
ments. As argued above in connection with equilibrium models, even small
departures from a model's assumptions may profoundly alter major eco-
logical phenomenon such as community stability I competition, and ecosystem
evolution. Theoretical ecologists in increasing numbers (e.g., den Boer, 1980;
Giesel, 1974; Wangersky, 1978; Wiens, 1977) are arriving at a viewpoint
similar to that expressed by Neill: Vfhe nearly complete dependence of
theoretical community ecology on a limited number of simple models [such
as the Latka-Volterra competition equations] has resulted in the development
176
of subsequent generations of deceptively simple constructs that may often
confuse our understanding of the underlying biology rather than inlprove
it' (1974, p. 399).
Acknowledgments
Special thanks are due to Michael Turvey and Claudia Carello for their part
in our discussions of some of the problems addressed in this paper. I am also
grateful to them, and to Bill Mace, Tim Johnston, Ed Reed, Bill Warren, and
an anonymous reviewer for their comments on earlier drafts of this paper.
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