The Niche Concept Revisited: Mechanistic Models and Community Context

Author(s): Matthew A. Leibold

Source: Ecology, Vol. 76, No. 5 (Jul., 1995), pp. 1371-1382
Published by: Ecological Society of America
Stable URL: http://www.jstor.org/stable/1938141 .
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 Editor's
Note
Ecology's love-hate relationship with the niche concept has been long and not
especially pretty. Whereas some ecologists continue to see value in the concept,
others have despaired of ever finding an expression that is both general and non-
circular. Part of the problem, originating early in our science, has been that different
investigators have meant different things by "niche." Here, Leibold uses resource-
Emphasizing based competition theory to produce a single construct that accommodates the
earlier differences in meaning, while at the same time permitting measurement
new ideas and experimentation. Although attempts to define the niche of a species may still
to prove fruitless, Leibold's contribution is to suggest that it is nevertheless possible
to delineate niche parameters relevant to a variety of ecological interactions.
stimulate
research Nelson G. Hairson, Jr.
inecology
Ecology, 76(5), 1995, pp. 1371-1382
© 1995 by the Ecological Society of America

THE
NICHE
CONCEPT
REVISITED:
MECHANISTIC
MODELS
ANDCOMMUNITY
CONTEXT
MathewA. Leibold
Department of EcologyandEvolution
of Chicago,
University 1101E.57thStreet
Chicago,Illinois
60637USA

Abstract
The niche concept is a central organizing aspect of modern ecology. Although
its history has often been reviewed, the structure of the concept and its connection
to advances in ecological theory has received less recent attention. I review the
niche concept using "mechanistic" models of community theory to identify two
distinct components. One describes the environmental requirements of organisms
and the other describes the per capita impact of organisms on the environment. I
argue that these correspond to significant differences between Grinnell's and El-
ton's concepts distinct from the previously discussed "habitat" vs. "functional"
dichotomy. I illustrate the distinction between the requirement and impact com-
ponents of the niche using models of resource competition and of keystone pred-
ators, and I discuss "Gause's axiom" and conventional "niche theory" in the
context of these two distinct niche components. I suggest that the niche concept
be elucidated by explicit reference to these two distinct components; the "impact"
niche (corresponding to Elton's concept) describing instantaneous per-capita ef-
fects of species on the environment, and the "requirement" niche describing the
response of species to the environment (corresponding to Hutchinson's definition).
This approach connects conventional niche theory with "mechanistic" individual-
based ecological models and can help provide a more modern context for the niche
concept.
Key words: competition; niche; predation; species impact; species requirement; species
role; zero net growth isoclines.

Introduction
Although most textbooks emphasize the use of the niche concept in community
ecology (e.g., Ricklefs 1979, Begon et al. 1990, Pianka 1994), the concept (or a
near synonym) is used by ecologists working at most levels of ecological orga--
nization. Thus, physiological ecologists often work to identify environmental con-

Manuscriptreceived 9 June 1994; revised 7 November 1994; accepted 24 November

1994.

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1372 MATHEWA. LEIBOLD
ditions (including factors other than resources) that
affect an organism's performance (and implicitly,
some component of its fitness). Similarly, much of
population biology is concerned with identifying lim-
iting factors of the environment that can alter the dy-
namics of populations (whether density dependent or
not). Biogeographers consider how environmental
conditions can constrain the distributions of taxa, and
ecosystems ecologists seek ways of describing how
the functional traits of taxa (whether species or func-
tional groups) alter ecosystem processes or structure.
The niche concept is not always used explicitly in
theories at all these levels of organization, but it pro-
vides an important, if sometimes implicit, connection
between these disparate fields of ecology that justifies
its status as a fundamental ecological concept (Cher-
rett 1989, Real and Brown 1991). Despite its strong
synthetic role and its crucial importance in community
theory, the niche concept remains unclear: "most
[ecologists] would agree that niche is a central concept
of ecology, even though we do not know exactly what
it means" (Real and Levin 1991).
In this paper, I re-examine the niche concept using
"mechanistic" models of species interactions. I sug-
gest that there are two closely related but potentially
distinct types of trade-offs that affect species inter-
actions related to the niche and I argue that each type
of trade-off is implicitly related to the two different
niche concepts. The first corresponds to Grinnell's
(1917) and Hutchinson's (1957) concepts concerning
environmental requirements of species, whereas the
second is more closely related to Elton's (1927) and
MacArthur and Levins' (1967) concepts concerning
the short-term impacts of species on resource use. I
conclude with the suggestion that the niche concept
be modified to combine these two narrower concepts
and I employ this broader concept to illustrate the link
between current "mechanistic" community theory and
conventional "niche theory."
A Historical
Evaluation
Schoener (1989), Griesemer (1992), and Colwell
(1992) have provided recent historical overviews of
the niche concept that contrast with earlier views of
Allee et al. (1949), Whittaker et al. (1973), and Hutch-
inson (1978). These discussions focus largely on dis-
tinguishing between "habitat" and "functional" as-
pects of the niche and on the importance given to the
relation of different species' niches to each other (see
also James et al.1984). In the discussion below I will
focus instead on the distinction between environmen-
tal requirements and environmental impacts of spe-
cies to highlight the dichotomy between the responses
of organisms to the environment and their effects upon
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Ecology, Vol. 76, No. 5
it (see Goldberg 1990 for a related analysis of aspects
of competition in plants). My conclusions are more
similar to those of Whittaker et al. (1973), who seem
to have anticipated many of the discomforts with the
niche concept that motivated my analysis, but who did
not focus on an explicit distinction between a species'
requirements and its impacts.
When Grinnell (1917) described the niches of or-
ganisms, he emphasized the multitude of environmen-
tal requirements of species and ignored, or only men-
tioned in passing, the potential impacts on other or-
ganisms. Grinnell included all of the commonly listed
environmental "limiting factors" in his formal de-
scriptions of species' niches. These included micro-
habitats, abiotic factors, resources, and predators, and
he emphasized the combination of physiological and
behavioral adaptations that allow species to respond
to such factors. In an era when field experiments were
uncommon, he explicitly tried to identify conditions
that allowed a given species to exist by examining the
union set of conditions in all the environments where
it was found. This approach seems entirely consistent
with other population-based approaches, such as An-
drewartha and Birch's (1954, 1984) "ecological web"
and Shelford's "environmental complex" (1913), and
it illustrates the great importance that ecologists have
placed on describing such factors at least since the
time of Darwin (1859). These concepts appear syn-
onymous with Hutchinson's (1957, 1978) "fundamen-
tal niche," which he defined as a multi-dimensional
"hyper-volume" describing the conditions where an
organism's expected absolute fitness is at least zero,
in a conceptual space whose axes include all of the
environmental variables affecting that species. Hutch-
inson argued that this definition would "completely
define its ecological properties" (Hutchinson 1957:
416), though it is easy to see ways in which this might
not be so (e.g., the incidental impacts of large grazers
on plant soil structure by compaction).
Elton used the niche concept in a distinctly different
way to describe the effect that a given species has on
the environment, sometimes including abiotic factors.
Although Elton (1927) discussed "limiting factors"
such as those described by Shelford (1913) and Grin-
nell (1917) at length, devoting an entire chapter (Chap-
ter 2) of his book to them, that chapter does not discuss
the niche concept. Instead, Elton devotes a section of
a separate chapter on community relations (Chapter 3)
to the niche. There, he describes the niche as an in-
creasingly fine description of the "role" of a species
in the community, beginning with its trophic level. It
is hard to know if the primary importance he placed
on trophic level reflected Elton's view that this was
one of the more important ways to describe the re-
July 1995 REVISIONINGTHE NICHE
source requirements of a species, or if he viewed it as
one of the more important ways to describe which
aspects of the ecosystem it was most likely to affect.
Elton's discussion of limiting factors and species re-
quirements in his Chapter 2 shows that he appreciated
the tremendous importance of abiotic factors on spe-
cies, but his explicit negation of most of them in de-
fining the niche of a species (Elton 1927: 64) suggests
a focus on what he calls "roles" or "what they are
doing." Though Elton may have meant to include spe-
cies requirements in his definition of the niche (thus
anticipating much of the following argument), I be-
lieve Elton's focus was on the impact of a species on
the ecosystem for the following reasons:
1) The niche was "defined to a large extent by [a
species'] size and food habits." Elton gave tremen-
dous significance to body size as a component of the
niche, not because it influenced requirements (as might
have been suggested by reference to the effects of body
size on thermoregulation, for example) but because
body size influenced what resources could be con-
sumed and what predators could be dangerous for a
given organism.
2) Elton's niche concept was intimately related to
the trophic pyramid of numbers, food chains, and food
cycles, and patterns of relative body size between
predators and prey describing interrelations among
species. Elton specifically notes that niches are im-
portant because they "enable us to see how very dif-
ferent animal communities may resemble each other
in the essentials of organization." This does not seem
related to an interest in the general set of all factors
that can allow a species to exist in different environ-
ments. It rather seems to reflect an interest in the sim-
ilarity of impacts of species across community types
rather than similarities in their requirements. Elton
described species with very distinct requirements
(e.g., Arctic foxes and African hyenas) as having the
"same" niche because they ate similar things. It is
hard to imagine Grinnell saying the same thing, though
Grinnell did claim identity of niches between species
like jerboas and kangaroo rats that presumably had
very similar requirements as well as similar re-
sources in different parts of the world.
3) Elton (1927: 64) specifically states that most
niche relations do not involve abiotic factors. How-
ever, he later mentions examples wherein species have
impacts on abiotic factors: the roles of earthworms
and other burrowing organisms (including land-crabs)
in affecting soil structure, and the roles of Sand Mar-
tins and Bee-eaters in digging out sand banks. He also
discusses the role of holothurians in converting corals
to "mud," which illustrates that "the coral-eating
niche has a geological significance" (Elton 1927: 68).
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1373
Thus Elton's view of the niche was much more ori-
ented toward describing how an organism affected the
environment (primarily by consuming resources and
serving as resources for higher trophic levels) than in
describing what environmental factors affected the or-
ganism. I argue below that this view corresponds more
strongly with MacArthur and Levins' use of the con-
cept describing resource consumption than with either
Hutchinson's definition or Grinnell's concept. Deter-
mining the difference between the two versions of the
concept is difficult because resources must be con-
sumed if they are needed to satisfy any "require-
ments." Furthermore, on a long-term quasi-equilib-
rium level, the full impact of a species on the envi-
ronment depends on a complex interaction of both of
these niche concepts with features of the environment
as discussed below.
Hutchinson contributed to the confusion by using
the concept ambiguously even in his seminal "Con-
cluding Remarks" paper (Hutchinson 1957; see also
Hutchinson 1978). Although Hutchinson used a defi-
nition of the niche explicitly focused on requirements,
his use of the niche in discussing competition was
much more compatible with a concept related to im-
pacts and roles of species. In his definition of the fun-
damental niche, Hutchinson specifically included
quantitative axes for every factor that could influence
fitness. Thus there could be an axis quantifying tem-
perature, the concentration of hydrogen ions (pH), etc.
However when discussing resources, Hutchinson il-
lustrated his point in a qualitatively different way.
Instead of having an axis that described the levels or
concentration of each potential resource (a natural ex-
tension of his use of the concept for abiotic factors),
he used axes that served as quantitative descriptions
of resources (e.g., food particle size, or distribution
along a habitat gradient). This difference is important
because information about resource levels was sacri-
ficed in order to reduce the dimensionality of the prob-
lem by using a single axis to describe (actually to serve
as an "index" for) a potentially large number of dif-
ferent resources. As discussed by Schoener (1989) and
others (see Giller 1984), Hutchinson's approach cre-
ated problems in understanding the association be-
tween resources and fitness and could not deal with
resources that could not be described by simple quan-
titative axes. Considering that, without knowing their
density, it is impossible to determine if resources are
adequate to allow survival and reproduction, there is
a severe deviation in his use of the concept from his
original definition, an observation that is commonly
made even by undergraduate students exposed to
Hutchinson's paper (M. Leibold and E. Simms, per-
sonal observation). The analysis described below,
 1374 MATHEWA. LEIBOLD
O )
:I O
° (O
~a>
E Arthur (1970, 1972), Maguire (1973), and Tilman
R*
Resource Density
FIG. 1. Resource-dependent (solid line, labeled b) and
resource-independent (dashed line, labeled d) fitness com-
ponents. Resource-dependent fitness components are thought
to affect individual growth and reproduction and should be
positive, whereas resource-independent components are
thought to depend on individual respiration and senescence
and should be negative. An equilibrium occurs when the
absolute values of the two components are equal and deter-
mines the minimum resource requirements of the organism
to just replace itself; it is associated with an absolute fitness
of zero when resource levels are at R*.
however, shows that niche relations among species can
be analyzed with an approach entirely consistent with
Hutchinson's definition without resort to ignoring re-
source-level effects on fitness.
In taking an approach different than Hutchinson's,
Maguire (1973) tried to re-describe the niche in terms
of resource levels in a fashion identical to that de-
scribed below. Hutchinson (1978) seems to have
agreed with the compatibility of Maguire's approach
with his niche concept (citing it as "doubtless the most
important contribution to the concept of the niche
since the death of Robert MacArthur"), but did not
seem to recognize the significance of the distinct use
of resource axes in the two approaches. The theoretical
basis for this distinction initiated by Maguire (1973)
combined with more recent theoretical work provides
a framework for making this distinction explicit.
A MechanisticAnalysisof ResourceNiche Relations
Following Hutchinson's work it was widely rec-
ognized (see Begon et al. 1990, Abrams 1992) that
there is a distinction between resources (defined as
depletable, potentially limiting, factors in the envi-
ronment) and other environmental factors that limit
populations (e.g., temperature, or disturbances). Ap-
proaches that explicitly consider the process of re-
source depletion have allowed ecological theory to
move away from phenomenological models based on
the highly "abstracted" (sensu Schaffer 1981) Lotka-
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Ecology,Vol. 76, No. 5
Volterra models to more mechanistic models (e.g.,
MacArthur 1972, Schoener 1974, Tilman 1982). These
explicit resource competition models provide an im-
portant insight into the distinctions that I believe dis-
tinguish the Grinnell/Hutchinson niche and the Elton/
MacArthur-Levins niche. The essential elements of the
competition model are described in detail by Mac-
(1982) and summarized and compared by Naeem and
Colwell (1991). Here I give an abbreviated description
to illustrate the distinction between aspects of the bi-
ology of organisms that relate to their requirements
and those that relate to their impacts on resources.
The interaction between resources and consumers
depends in part on how resources affect fitness com-
ponents of the consumers. This can be illustrated as
shown in Fig. 1 by separating a resource-dependent
component (hereafter referred to as the population
growth rate, b) and a resource-independent component
(hereafter referred to as the loss rate, d), recognizing
that each of these components involves both repro-
duction and survival probabilities related to produc-
tion and respiration processes. If resource levels are
high enough for the growth rate to exceed the loss
rate, the population will increase; if the resource levels
lead to a growth rate lower than the loss rate, the
population will decline. There is a set (only one point
in Fig. 1, labeled R*) of conditions where the growth
rate equals the loss rate.
Following Maguire (1973, see also Hutchinson
1978, Tilman 1982), assuming that other factors are
held constant, and focusing on a pair of resources
R1
FIG. 2. Resource-dependent fitness of a consumer feed-
ing substitutably on two resources. Resource densities are
shown on the two axes. The line shows the combination of
resource densities that allow the consumer to have an ab-
solute fitness of zero (labeled ZNGI) and delimits conditions
where the consumers fitness is less than zero (open area) and
greater than zero (hatched area). Using Hutchinson's formal
definition, the consumer's niche is described by the hatched
area.
July 1995 REVISIONINGTHE NICHE
(whose densities are labeled R, and R: in Fig. 2), one
can describe three sets of conditions for any consumer
that depends on these two resources. If resources are
too low (the clear area in Fig. 2) the consumer's den-
sity-independent fitness loss component (d) will ex-
ceed the resource-density-dependent growth rate (b).
By Hutchinson's definition such conditions are "out-
side" the consumer's niche. In contrast, resource lev-
els may be high enough (the hatched area in Fig. 2)
that the growth rate exceeds the loss rate and the pop-
ulation can increase. By Hutchinson's definition, such
conditions are "inside" the consumer's niche. A third
set of conditions obtains when both fitness components
are equal, since the organism will have an expected
fitness of zero and will neither increase nor decrease.
This set of conditions determines the "zero net growth
isocline" or "ZNGI" (Tilman 1982) of the organism
and serves to decribe the "boundary conditions" that
delineate the organism's Hutchinsonian niche. Though
the derivation of the ZNGI does not depend on pop-
ulation equilibrium assumptions, MacArthur (1972),
Maguire (1973), and Tilman (1982) all point out that
long-term numerical responses will tend to lead to
conditions on the "ZNGI" as density-dependent pro-
cesses occur if the consumers deplete the resources.
However, resource-consumer relations described by
the ZNGI line are not fundamentally different from
relations involving any other (non-resource) factor,
and the relationship between resource levels and fit-
ness does not explicitly describe resource consump-
tion. It is possible to have two species with identical
ZNGIs but whose resource consumption rates are very
different (see below) and vice versa.
Resource consumption can be described by a dif-
ferent set of relations illustrated by vectors relative to
R, and R, as shown in Fig. 3 (see MacArthur 1972,
Tilman 1982). The short-term impacts of a consumer
on the two resources depend on the rates at which they
consume each of the resources. The magnitude of sep-
arate vectors parallel to each resource axis can de-
scribe the expected rates at which each individual con-
sumer consumes each resource. This can, and usually
will, depend in a complex way on the abundances
the resources and will be affected by the functional
responses of organisms. However, at any given set of
resource densities, the net expected effect of each con-
sumer can be described by adding the two orthogonal
vectors to define an "impact vector" (Tilman refers
to this as the "consumption vector" but I wish to
generalize the concept below and to emphasize its role
in discerning the effect of the consumer on the re-
sources). An important point to make is that such vec-
tors do not necessarily depend on consumer density,
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1375
per capitafeeding rate
on R,
/ percapita
c\j
'foorlinn
1W1u 'Y
Cc rate on R2
C;
Impact
Vector
R1
FIG. 3. Per capita effects of a consumer on two substi-
tutable resources. Axes as in Fig. 2. Each individual con-
sumer has an expected impact on each resource described
by the vectors with thin lines. The magnitude of the vector
is proportional to the feeding rate of the consumer on each
resource. The total impact of the consumer on the two re-
sources is described by the sum of these components, shown
by the thick arrow labeled C.
and that their derivation does not depend on any as-
sumptions about population equilibria.
The relationship between impact vectors and ZNGIs
depends strongly on the efficiency of the consumer in
converting resources to per capita growth. Thus, two
consumers might have similar ZNGIs despite having
very different impact vectors if they also differ in the
relative efficiency with which they convert each re-
source to growth. In the simplest linear model with
substitutive resources (sensu Tilman 1982) comparing
two species with identical loss rates and identical mag-
nitude (but not slope) of impact vectors, a pair of
species would have the same ZNGI if the ratio of their
consumption rates is equal to the inverse of the ratio
of their efficiencies on the two resources. The long-
term population-level impact of the organism depends
on an interaction between the numerical response of
the organism (which depends on its ZNGI) and its per
capita impact vector at equilibrium.
Similarly, the basic principles described for com-
petition for two resources can also be used to examine
of interaction involving a single resource and a predator
on an intermediate consumer species (Holt et al. 1994,
Leibold, in press). By separating fitness components
into those that depend on resources and predators from
those that are density independent, we can plot the
combination of resource and predator densities that
result in an expected fitness <0 (population declines;
the open area of Fig. 4), >0 (population growth; the
hatched area in Fig. 4) and fitness =0 (the ZNGI line
shown in Fig. 4). These conditions describe when re-
source levels are high enough to mitigate the negative
1376 MATHEWA. LEIBOLD
(I)
0)
Resource Density
FIG. 4. Effects of a predator and a resource on a species
with an intermediate position in a food chain. Notation as
in Fig. 2. Using Hutchinson's formal definition, the organ-
ism's niche is described by the hatched area in the graph.
effects (either involving direct mortality or reduced
performance of the species due to its facultative re-
sponses to predators) of predators on the expected fit-
ness of the organism. Again, the hatched area satisfies
Hutchinson's definition of the niche but does not di-
rectly describe how organisms affect their prey (via
consumption) nor their predators (via serving as food).
The ZNGIs describe the environmental requirements
(and constraints) but do not describe the ecological
impacts on the predator and resource except those that
are eventually expected to occur via long-term nu-
merical responses if the species can modify predator
densities, resource levels, or both.
As before, the short-term ecological impact of the
species can be described using vector notation (Lei-
bold, in press) to partition the expected per capita
effects of the organism into the effect on its resource
and the effect on its predator. The vector component
describing the organism's impact on its resource is
identical to the one used above to describe competition
for two resources. The impact on predators is de-
scribed by a vector parallel to the predator axis whose
magnitude is equal to the product of the death rate
imposed by the predator on the organism and the pre-
dator's conversion efficiency (i.e., the per capita con-
tribution of the organism to the predator's population
growth rate). The net per capita effect of the organism
on the environment (the resource and the predator) is
described by the sum of the component vectors, which
defines an "impact vector" (labeled C in Fig. 5). In
contrast to the model for two resources where impact
vectors point toward the lower left (negative impacts
on both resources), here impact vectors will generally
point to the upper left (positive impact on predators
and negative ones on resources). Again, the deriva-
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Ecology,Vol. 76, No. 5
tions of the ZNGI and the impact vector do not depend
on population equilibrium assumptions.
As before, species can have similar ZNGIs yet have
distinct impact vectors. For example, consider a case
where two species have identical minimum resource
requirements in the absence of predators and are equal-
ly assimilated by. the predator and have linear func-
tional responses. Such species will have impact vec-
tors whose slope is their mortality risk to predators
divided by the feeding rates on their own resource.
Species could have identical ZNGIs despite having
different such vectors if the ratio of their relative as-
similation efficiencies on their own food is equal to
the ratio of the slopes of their impact vectors.
Again, though ZNGIs and impact vectors are related
to each other, they describe two potentially distinct
aspects of the relationship of an organism with exter-
nal environmental factors. ZNGIs correspond closely
to Hutchinson's niche concept and to Grinnell's de-
scription of biotic factors that affect the fitness of or-
ganisms (including predators), whereas impact vectors
reflect the roles that species play in the community by
culling resources and contributing to the support of
higher trophic levels as discussed by Elton. In this
case the distinction is even more striking by its rele-
vance to food chains and the "pyramid of numbers."
Coexistenceandthe Niche
Regardless of which concept is involved, the niche
has always been closely associated with "Gause's ax-
iom," that two species cannot coexist if they share a
single niche (Gause 1934). The theoretical models de-
> per capita
.)_
C
C; mortalityrisk
VImpact x predator
QC]~ Vector assimilation
per capita
Q. feedingrate -_
on resource
Resource Density
FIG.5. Impactof a species with an intermediateposition
in a food chain on its resourceand on its predator.Notation
is similar to Fig. 3 except that the impact of the organism
on its predator(the verticalcomponentarrow)is proportional
to the mortalityrate experienced by the organism due to
predation, multiplied by the assimilation efficiency (rate of
conversion of prey to growth and reproduction) of the pred-
ator. The net per capita impact vector is the sum of the
components shown with a thick arrow labeled C.
July 1995 REVISIONINGTHE NICHE
scribed above explicitly examine conditions for co-
existence, and they identify two different criteria re-
lated to the niche. Furthermore, both aspects of the
niche that can be identified by using these mechanistic
models are determined by the expected (i.e., average)
parameters of individuals without reference to popu-
lation equilibria (though of course the prediction itself
is only expected to hold if populations have come to
near-equilibrium conditions): First the expected en-
vironmental requirements for an individual to com-
plete its life cycle and replace itself, and second, the
expected per capita impact of individuals on other or-
ganisms.
I argue that these two different criteria are identified
with two different concepts of the niche; one associ-
ated with Grinnell (1917) and Hutchinson (1957,
1978), and the other associated with Elton (1927) and
MacArthur and Levins (1967). In models of compe-
tition for two resources (e.g., MacArthur 1972, Tilman
1982) and in the model involving interactions among
two species that share a common resource and predator
(Holt et al. 1994), coexistence of two species requires
that their ZNGIs cross. These aspects of the models
describe conditions for the EXISTENCE of an equi-
librium point allowing coexistence of species. Ignor-
ing the "infinitely small" possibility that different
species will have identical ZNGIs, this suggests that
species must have different ZNGIs to coexist and that
these differences must further be of the type where
each species has higher fitness under some environ-
mental conditions. For example, in the two-resource
model each resource must benefit one species more
than the other. Likewise, in the keystone-predator
model, one species must be less affected by the pred-
ator and the other must be a better resource exploiter
(sensu Tilman 1982) in the absence of predators. The
existence of this equilibrium point does not depend
on the impact vectors (except indirectly due to shared
parameters common to both ZNGIs and impact vec-
tors).
Given the existence of an equilibrium point as de-
scribed above, however, the stability of the point de-
pends critically on the impact vectors (Tilman 1982,
Leibold, in press). In the case of competition for two
resources, stability requires that each species con-
sumes proportionately more of the resource that most
strongly limits its growth (Tilman 1982). In the case
of competition mediated by a keystone predator, sta-
bility requires that the species most strongly affected
by the predator (i.e., the "resource-exploitation spe-
cialist") contribute proportionately more to the pre-
dator's growth relative to its feeding rate (M. A. Lei-
bold, unpublished manuscript).
Thus the joint criteria for a stable equilibrium re-
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1377
quires a linked set of trade-offs between species; one
set must reflect a trade-off in the requirements of spe-
cies, and the other must reflect a corresponding trade-
off in the impacts of species. Of course, because both
aspects are affected by some common parameters (di-
rect mortality and feeding rates), both conditions for
a stable equilibrium can be satisfied by trade-offs in-
volving a single variable (relative feeding rates in the
case of competition for two resources and the "impact
ratio" described above for the keystone predation
model). However, an important point is that trade-offs
need not necessarily involve a single variable because
such trade-offs could also involve assimilation and
loss rate parameters.
The two examples discussed above (competition for
two resources and competition mediated by a keystone
predator) are only two of a wide array of reciprocally
negative species interactions that can result from many
different mechanisms. These examples only illustrate
some of the points that characterized Grinnell's and
Elton's views about niches and serve as a starting point
for a similar analysis of other mechanisms. To date
models of species interactions such as those described
above are best understood in closed (i.e., self-sustain-
ing local populations), homogenous (no patchiness or
disturbance) situations. They can, however, serve as
the basis for more complex situations, as illustrated
by the work on environmental heterogeneity by Levins
(1979), Chesson (1986), and Tilman (1982; see also
Naeem and Colwell 1991), which illustrate the in-
creased dimensionality of the problem when compared
to the simple cases described above.
As has been often noted (e.g., Levins 1968, Tilman
1986), virtually all of the mechanisms proposed to
explain long-term coexistence of species can be related
to trade-offs in some components of their biology or
of their "niches" (Chesson 1991). It is not yet clear
how many different mechanistic models of species in-
teractions can be analyzed in a way that separates the
conditions for existence from those that determine sta-
bility of putative equilibria into the respective com-
ponents of the niche concept involving requirements
and impacts.
Short-term
vs. Long-term of Species
Impacts
One of the attractive features of the "mechanistic"
approach is that biological parameters are described
at the individual level (or at the level of the entire life
history of individuals). Separating "requirement" and
"per capita impact" components of the niche are en-
tirely consistent with this "individual-based" ap-
proach, and the concepts do not depend on making
equilibrium assumptions about the component popu-
lations of interacting species (including resources,
 1378 MATHEWA. LEIBOLD
competitors or predators). However, on a long-term
basis the total impacts of a species depend on its pop-
ulation size, and the most predictable aspect of such
long-term impacts are those that occur at the popu-
lation equilibrium. In both of the models described
above (involving competition for shared resources and
predator-mediated competition) the long-term, equi-
librium, impact of species depends more closely on
their requirements than on their per-capita impacts (see
Tilman 1982, Holt et al. 1994). Thus Tilman's (1982)
conclusion that species with the highest competitive
ability should be those with the lowest resource re-
quirements (the "R* rule"), and Holt et al.'s (1994)
analysis of apparent-competition ability (with an anal-
ogous "P*" rule) illustrates that on a population level
that assumes an equilibrium, impact vectors do not
play as important a role as do ZNGIs. However, a focus
on describing relative competitive ability that ignores
impact vectors cannot elucidate the important roles
that species play in the dynamic aspects of community
stability via their relative abilities to affect the envi-
ronment nor can they describe the roles that species
play in regulating turnover-rates of their resources or
in supporting populations of their predators (see Til-
man 1982, Holt et al. 1994). Furthermore, an approach
focused on long-term impacts (e.g., using ZNGIs only
to predict competitive ability) that ignores impact vec-
tors cannot help make predictions about which species
can coexist since it cannot predict which combinations
allow for stable coexistence.
Relation
of theMechanistic to Conventional
Approach
"NicheTheory"
Which of the two aspects of the regulation of co-
existence is included in conventional niche theoretical
models? Modern "Niche Theory" focuses on exam-
ining niche relations with respect to resource con-
sumption (see Giller 1984, for a recent review). De-
spite the strong personal connection between Hutch-
inson and MacArthur as they developed current niche
theory, and acknowledging Hutchinson's ambiguous
use of the concept with respect to his formal definition,
I argue that modern niche theory most closely relates
to impact vectors and has little to do with resource
requirements (despite the observation that some def-
initions of competitive ability depend more on require-
ments than on per capita impacts as described above!).
Modern niche theory owes much of its development
to the idea, proposed by MacArthur and Levins (1967),
showing that Lotka-Volterra competition coefficients
could be equated with diet overlap of competitors.
Although subsequent work (see Giller 1984), especial-
ly by MacArthur (1970, 1972), Schoener (1974), and
Abrams (1975), has shown that this is not strictly true
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Ecology, Vol. 76, No. 5
under more complex situations, the general result has
continued to be used as a practical "rule of thumb"
(Chesson 1990). There has subsequently developed a
large literature deriving quantitative metrics of "niche
overlap" and "niche breadth" (see Petraitis 1979 for
a review) and studying resource use overlap in natural
communities (e.g., Cody 1968, Pianka 1973). There
are two ways of looking at this body of work to see
that they describe aspects of niche relations related to
environmental impacts rather than to environmental
requirements.
First, as shown by Vandemeer (1975), Lotka-Vol-
terra competition coefficients are not sufficient to de-
termine whether two species will coexist. Instead they
determine whether the equilibrium point will be stable
given that it exists (with both species having positive
densities). Thus meeting the conditions of "permis-
sible" niche overlap is equivalent to determining the
stability of possible equilibrium points if MacArthur
and Levins' (1967) isomorphism between overlap and
competition strength applies. Vandermeer's result
shows that permissible niche overlap is necessary for
a stable equilibrium but does not guarantee the exis-
tence of such an equilibrium point (see also Persson
1985).
More to the point, Petraitis (1989) has shown that
the cosine of the difference between the angles of im-
pact vectors of two species in Tilman's (1982) com-
petition model is equivalent to one of the specific mea-
sures of resource overlap based on the "conventional"
niche model (Pianka 1973). As discussed above, im-
pact vectors need not have a direct relation with re-
source requirements, because they are based on use
without respect to their effects on fitness. Rather, con-
clusions about coexistence based on differences be-
tween diets (conventional niche theory) are equivalent
to differences in per capita impacts (graphical mech-
anistic models) on resources. This result shows that
the overlap indices of conventional niche theory do
not describe the expected long-term effect of organ-
isms on the densities of their resources (i.e., their ef-
fects on equilibrium resource densities as modeled by
the intersection of ZNGIs using Tilman's terminolo-
gy), and thus do not fully describe the process of in-
terspecific competition but only describe the short-
term relations in relative impacts (impact vectors
only). Such confusion about competitive abilities and
the importance of short-term vs. long-term effects un-
derlies a number of current debates in ecology (e.g.,
disagreements between Tilman's and Grime's views of
"competitive ability," see Grace 1990, and Goldberg
1990 for recent reviews).
What is Hutchinson's "realized niche"? Hutchinson
(1957) introduced the "realized niche" as a concept
July 1995 REVISIONINGTHE NICHE
SI
FI(l. 6. Environmental conditions, dependent on the Sup-
ply Point (equivalent to equilibrium resource densities in the
absence of the consumers, described by the axes labeled S,
and S,) that determine the outcome of competition between
two species competing for substitutable resources. The solid
lines show the zero-net-growth isoclines of the two consum-
ers [labeled ZNGI (a) and ZNGI (b)j. The outcome depends
on their impact vectors (labeled C(a) and C(b)] as illustrated
by the extension of those shown as dashed lines. If maximum
resource densities in the absence of consumers occur in Re-
gion I, neither species can exist. In Region II, only species
h can exist, whereas only species a can exist in Region VI.
Though both species can exist in the absence of interspecific
competition in the remaining regions, species a is excluded
from Region V and species h is excluded from Region III
by competition. The two species coexist only if the Supply
Point is in Region IV. See Tilman (1982) for a detailed dis-
cussion. Petraitis (1989) has shown that the cosine of the
angle between the two impact vectors IC(a) and C(h)l is
identical to Pianka's (1973) measure of niche overlap.
to identify those types of niche axes that were influ-
enced by competitors (i.e., primarily resources) and to
describe how interspecific competition acted to limit
populations. The notion was to determine how an en-
vironment satisfied the requirements of an organism
before (the "fundamental niche" concept) and after
(the "realized niche" concept) accounting for the ef-
fects of competitors. Further, Hutchinson suggested
that environmental conditions describing overlap in
the fundamental niches of different species could be
partitioned between them to quantify the realized
niche. As was pointed out by Schoener (1989), there
were a number of conceptual problems associated with
Hutchinson's idea of such partitioning. However,
mechanistic models can be used to illustrate the basic
idea.
Using Tilman's (1982) formulation of resource com-
petition for substitutable resources we can imagine
defining species niches on the basis of the Supply Point
of the resources (i.e., resource densities in the absence
of consumers, before accounting for the effects of
competitors) as shown in Fig. 6. The only difference
in the notion of the "niche" between this figure and
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1379
Fig. 2 discussed above is the definition of the axes;
here the axes are supply points (what resource levels
would be if consumers were absent), whereas the axes
in Fig. 2 are resource densities (actual densities ex-
perienced by the consumers at a given point in time),
as discussed extensively by Tilman (1982). Different
possible environments are classified as having supply
points falling in various parts of the graph labeled with
Roman numerals. As discussed by Tilman (1982), spe-
cies are excluded from some such environments by
interspecific competition (In Fig. 6, species a is ex-
cluded from environments whose supply point lies in
region III, and species b is excluded from those found
in region V). Further, species can coexist only under
conditions, determined by their consumption (i.e., im-
pact) vectors, whose supply points are found in region
IV. This separation of environmental conditions that
determine where species exclude each other is a way
of partitioning of the "environmental space"; the rel-
ative densities of the species in the area allowing co-
existence in region IV could serve as weighting factors
to further describe the partitioning of the entire graph.
Again, partitioning the environmental space into the
different realized niches depends on the consumption
vectors much more than on the ZNGIs.
A confusing issue is that the overall competitive
effect of the competing species on resources is de-
scribed by the single point that describes the equilib-
rium outcome of the set of interactions (the intersec-
tion of the ZNGIs if the species coexist). Thus this
concept of the "realized niche" (defined as a subset
of the original supply points) cannot be quantified
from simple observations of natural communities be-
cause the position of the original supply points cannot
be reconstructed without conducting an experiment
(eliminating the consumers). However this concept
serves to show that the minimum resource require-
ments play a crucial role in determining which species
will win (equivalent to Tilman's "R* rule"; coexis-
tence depends on which species have the lowest min-
imum resource requirements; Tilman 1982). A similar
argument can be made to evaluate interactions among
species that share predators, and species that share
both resources and predators (see Holt et al. 1994).
In an apparent attempt to solve some of these prob-
lems with Hutchinson's niche concept and to make the
concept operational, MacArthur and Levins (1967) re-
defined both the fundamental niche and the realized
niche in terms of resource utilization (see also Schoe-
ner 1989). A multitude of other papers have since
followed that collectively form "conventional niche
theory" (reviewed by Giller 1984). Though much of
the distinction between "requirements" and "im-
pacts" outlined here was discussed by MacArthur
1380 MATHEWA. LEIBOLD
(1967, 1970, 1972), most "niche theory" has focused
on relative resource utilization without explicit ref-
erence to resource requirements (reviewed by Colwell
and Fuentes 1975 and by Giller 1984).
In "conventional niche theory," resources are made
available and are consumed by organisms in propor-
tions that depend on the occurrence (and density) of
other species. Thus competition describes the parti-
tioning of resources as they become available (a rate
described by resource renewal rather than a density
measure of availability), into different consumer pop-
ulations. The significance of Petraitis' (1989) result
described above is that it shows that this approach is
most relevant to the consumption ratios of species,
and that despite a superficial similarity to Hutchinson's
description of realized niches described above, it is
not necessarily closely related to minimum food re-
quirements.
Discussion
The significance of the recent focus on "mechanistic
models" (Price 1986, Schoener 1986, Tilman 1988)
is that they allow results of community-level models
to be defined on the basis of parameters that distin-
guish between the expected (i.e., average) properties
of individuals (e.g., feeding rates, assimilation effi-
ciency, and predator vulnerability) from those that are
properties of the environment (e.g., maximum re-
source levels, disturbance level, and abiotic stress). In
contrast, much conventional niche theory used a more
phenomenological approach (based on Lotka-Volterra
competition models) to evaluate to dynamic properties
(i.e., stability properties) of competition. The two ap-
proaches have sometimes appeared to be distinct and
somewhat disparate. As implied in the work of Mac-
Arthur (1970, 1972, see also Chesson 1990), I believe
a synthesis is possible based on the joint consideration
of Hutchinson's definition of the niche and Elton's
view of the concept as follows:
1) Environmental requirements do not necessarily
correspond to the per capita impacts of species on the
environment (i.e., there is not a necessary one-to-one
correspondence between ZNGIs and impact vectors).
2) The requirements of different species determine
whether coexistence is possible, and coexistence can
only occur if there is a trade-off in species' require-
ments. Such relationships also determine which spe-
cies combinations are most likely to coexist.
3) Whether coexistence will be achieved (because
it is stable) in a given environment depends on the
relationship among the impacts on the environment of
the species that have different requirements. Each spe-
cies must have a differential effect on the factor that
most limits its growth for coexistence to be stable.
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Ecology,Vol. 76, No. 5
4) Aspects of the niche related to environmental
requirements of species most closely correspond to
Grinnell's use (1917) and Hutchinson's (1957) defi-
nition of the concept as shown by Hutchinson's ac-
ceptance and discussion of Maguire's (1973) interpre-
tation.
5) Aspects of the niche related to environmental
impacts relate more closely to Elton's (1927) concept.
They also correspond strongly to MacArthur and Lev-
ins (1967) development of "conventional niche the-
ory" as shown by Petraitis' (1989) demonstration of
an isomorphism between niche overlap (and breadth)
measures and relations among Tilman's (1982) con-
sumption vectors.
6) A re-interpretation of the niche concept as the
union of these two niche concepts can allow greater
mechanistic insights into factors that determine the
outcome of species interactions as shown by the cor-
respondence in the analysis of niche relations in mod-
els of two-resource competition and of keystone-pred-
ator mediated competition for a single resource.
The niche concept has always been meant to be
general (and perhaps intractably so). Grinnell, Elton,
Hutchinson, and MacArthur and Levins all conceived
that it was meant to summarize either "many" or "un-
countable" aspects of the biology of organisms. Dis-
tinguishing between the two different views of the
niche concept clarifies it by identifying a different
theoretical consequence for each component (exis-
tence of an equilibrium vs. the stability of the equi-
librium). Further, it is clear that the two aspects are
closely (though not necessarily) linked in many situ-
ations by sharing common parameters; in the case of
trophic relations these are feeding rates and mortality
risks to predators. I suggest that the term "requirement
niche" be used to describe requirements (most closely
corresponding to Hutchinson's definition) and that the
term "impact niche" be used to describe the per capita
effects of species on their environments (correspond-
ing to Elton's "roles"). The two aspects combine to
form the "total niche" in its broadest sense. In much
"conventional niche theory," aspects related to im-
pacts of species have been emphasized as the ones that
regulate coexistence by affecting the stability prop-
erties of the interactions, but such considerations are
irrelevant if the conditions that permit coexistence (re-
gardless of the stability features of the equilibrium)
do not apply: Tilman's (1982, 1988) "R*" rule-of-
thumb must apply before the question of stability re-
lated to the "overlap" rule-of-thumb can even be con-
sidered.
My primary goal has been to highlight these results
with the hope of clarifying the niche concept and elu-
cidating its relationship to mechanistic models of com-
July 1995 REVISIONING THE NICHE
munity interactions. Clarifying this relationship also
allows a much closer linking of individual ecology
(involving physiology and behavior) to community
models without generating "abstracted parameters"
such as "competition coefficients" that are mixed
properties of populations, individuals, and environ-
ments (discussed by Shaffer 1981). Though similar
arguments related to resource competition were made
by MacArthur (1967, 1972), these have subsequently
received little attention in the context of "niche the-
ory" and have not been extended for use with other
types of species interactions (e.g., predator/prey in-
teractions). Such a mechanistic approach isf also more
compatible with the current emphasis on experimental
methods that combine ecological insights at the in-
dividual, population, and community levels (see Di-
amond 1986).
Use of the niche concept has experienced a notable
decline in recent years (Schoener 1989, Colwell 1992).
I believe that a major reason for this involves con-
fusion about its role in generating testable hypotheses
and its ability to synthesize basic ecological principles
from different levels of ecological organization (in-
dividuals, populations, communities, and ecosystems).
Chesson's (1991) call for a "Need for niches" on the
basis of trade-offs that can determine coexistence and
its stability, as described above, is an important way
to avoid sacrificing such an important concept to the
vicissitudes of increasing reductionism without com-
promising a strong mechanistic framework.
Acknowledgments
E. Simms, M. Wade, and E. Werner provided invaluable
comments on early versions of this manuscript. I also thank
J. Brown, D. Goldberg, R. Holt, M. McPeek, and T Schoener
for stimulating ideas and valuable criticisms.
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