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Opinion is intended to facilitate communication between reader and author and reader and
reader. Comments, viewpoints or suggestions arising from published papers are welcome.
Discussion and debate about important issues in ecology, e.g. theory or terminology, may
also be included. Contributions should be as precise as possible and references should be
kept to a minimum. A summary is not required.

How useful is the concept of habitat?

a critique

Sean C. Mitchell, 2-100 Highland Drive, Antigonish, NS, Canada B2G 1P6 (smitchel@stfx.ca).
The following argument regarding the quantitative value of the
habitat concept is presented for discussion purposes. The
discussion is not intended as a review of the habitat-population
literature (which is immense for any class of organism) and so does
not include an exhaustive appeal to authority. Instead, the
arguments are, largely, intended to stand or fall on their own
merit, and the literature is used selectively to demonstrate a point.

The concept of habitat is possibly the most fundamental, and unquestioned, paradigm in ecology. The
premise that the structure/condition of the physical
environment in which an individual organism, or group
of organisms, is found is critical to the population,
plays a central role in basic and applied ecology, where
the presumed relationship is used to interpret organism
distribution, evaluate population dynamics, and predict
abundance/biomass of organisms. In the applied field of
natural resources management the relationship is fundamental in assessing impacts of anthropogenic activities and predicting landscape level changes, restoring
perturbed systems, developing species-at-risk recovery
strategies, and inventory of species. Habitat is enshrined
in, and a fundamental component of, environmental
legislation, governing much of the assessment, mitigation or prosecution of anthropogenic activities. All of
these uses of the habitat concept rely heavily on the
presumption of some form of relationship between
population size (i.e. species abundance) and the physical
habitat in which the organism exists, and also that the
habitat in some manner limits that abundance. Such
relationships are intuitive and indeed in some cases are
observable (e.g. black footed ferret limited by prairie
dog habitat; piping plover by beach habitat for nesting,
etc.). Further, it is presumed that the measurement
(quantification) of various habitat and population
attributes will allow us to determine these relationships
and then to make decisions and take action with a
cause-and-effect relationship (e.g. providing or improving habitat will result in increased abundance). However, I propose for discussion that our present concept
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of habitat is flawed or not complete. I present the thesis

here that our present concept(s) of habitat are of value
only as qualitative descriptors of organism distribution,
and are of little value in quantitatively describing
organism abundance. As such, the concepts are of
very limited use in basic or applied ecology, or
quantitative management of populations. The following outlines the support for this thesis, beginning with
definition(s) and interpretation of habitat followed by
problems of measurement of pertinent variables, a case
study to demonstrate the point, and finally implications
if this thesis is correct.

Definitions/interpretation
For such a central-place concept in ecology the
definition of habitat must surely be among the least
rigorous of any in science. There does not exist any
authoritative definition; the most commonly recognized ones are Odums (1971) !/ the place where an
organism lives, or the place where one would go to
find it (p. 234) !/ and that of Whittaker et al. (1973)
who state that habitat is used for environment in its
physical and chemical aspects (p. 326), and that it is
usually conceived as the range of environments or
communities over which a species occurs (p. 328).
The critical point here, and for my argument, is that
habitat refers to the abiotic components of the
environment only. This is, as near as I can tell,
consistent with current usage (Hall et al. 1997,
Morrison and Hall 2002, Van Horne 2002). Both of
the above authorities also recognize that the term
habitat may be applied to an entire community in
addition to being specific to an organism. Thus, there
are three principle concepts of habitat, for each of
which I have provided my interpretation.
OIKOS 110:3 (2005)

Concept 1. Where an organism lives

This is a standing place or living space concept
(Looijen 1995) in which the presence of an organism is
correlated with environmental variables. This is clearly a
distributional concept only, it says nothing about how
habitat may interact with or affect abundance of
organisms. This particular concept may have use for
habitat specialists (e.g. coho salmon, spotted owl) but
has little meaning for those species ranging over wide
geographic areas of greatly varying conditions (e.g.
habitat generalists such as whitetail deer, coyote,
crow). As the presence of these latter organisms may
be correlated with almost all environments in North
America; this is not particularly instructive. Additionally, this concept does not provide us with any useful
information about the relationship of the organism and
its surroundings (e.g. the red squirrel is confined to
coniferous forests as habitat-that is a fine descriptor but
does nothing to explain the mechanisms controlling
squirrel abundance or range). Without providing some
understanding of how populations are controlled/limited
(e.g. food supply, predation, climate, etc.) this use of the
term habitat is, from a more than purely descriptive
perspective, not very useful. To use a theatrical analogy,
this concept of habitat merely provides the stage against
which the production takes place !/ but we are usually
much more interested in the plot and action of the play,
rather than the backdrop.

however, is that the limiting factor is not biological.

This is the crux of the issue !/ are organisms and
populations limited by abiotic or biotic factors? Of
course, this depends entirely upon the individual species
under consideration and will also depend upon its
abundance (at high abundance competition more likely
to have effect, at low abundance abiotic factors !/ the
old density dependence/independence debate). Thus,
this habitat concept may be more applicable under
some conditions (low abundance) than others (high
abundance).
My subsequent argument has been taken to task on
the grounds that this definition of habitat is too narrow
and workers today consider habitat to include biotic
aspects in addition to abiotic. My response is two-fold.
First, that merely demonstrates the ambiguity of what
we mean by habitat (a point made very well by Hall et al.
1997) in that some investigators limit it to abiotic only
and others include biotic aspects as well. Therefore, we
are measuring and including fundamentally different
variables and responses within the same nomenclature.
This can only lead to confusion and ambiguity. The
second part of my response is that if one is including
biotic factors, one is not evaluating habitat-abundance
relationships, but rather has strayed into the niche
concept and is misleadingly calling it habitat. The
distinction between the two is critical and in trying to
understand abundance and dynamics, the designation of
which concept is being employed is essential for clear
communication and understanding.

Concept 2. Environment in its physical and chemical

aspects

Concept 3. Community concept

This is intended as the set of environments meeting a

species ecological requirements and tolerances. That is,
the set of environments in which a species could, but
need not actually, live (Looijen 1995). Under this
concept, attempts are made to discriminate physical
and/or chemical variables (e.g. temperature, water availability, nutrients, etc.) affecting the survival or fitness
of individual organisms, or the success or dynamics of
the population. This approach may be viewed as
representing the subset of only abiotic dimensions of
the Hutchinsonian multidimensional niche (i.e. ignores
biotic variables such as competition, predation, allelopathy, etc.). This omission of biotic effects is generally
justified by the argument that while they are not part of
the habitat as defined, the physical and chemical
environment provides the conditions to meet these
challenges (e.g. climate to support food resources, space
to avoid predation and limit competition, etc.). Thus,
these biotic effects are implicitly included by this habitat
concept. This concept of the habitat being a function
solely of its abiotic variables is sensible and intuitive,
though reductionist. The single most critical assumption
linking the organism or population to this concept,

This idea may be summarized as the physical and

chemical environment in which a complex of plants
and animals exist (e.g. tall grass prairie habitat, running
water habitat). This is an extension to the larger
community of the relationship of the individual
organism to its environment, and as such it may be
considered an expansion of concept 2 in that it retains
the reliance on solely abiotic factors governing distribution and abundance. As I understand the concept, it is
intended to be purely descriptive; quantitative inferences
are not drawn regarding population abundance
or dynamics at this scale of organization. Therefore,
I will pass over this concept as it does not apply to the
developed thesis.
So, in summary, our current definition(s) of habitat
appear to provide fine descriptions regarding organism
presence and distribution. However, to either (a) understand mechanistic relationships between an organism
and its environment, or (b) predict changes in population abundance or dynamics due to changing conditions,
quantification of the relationship between the organism
and what we call its habitat is required. And this is where
significant problems arise.

OIKOS 110:3 (2005)

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Problems of measurement
I put forward that there are three principle problems in the
measurement of habitat and population attributes, and
the correlation of the two. These problems are of a
fundamental nature and I dont believe that they can
be overcome by technical solutions. The problems are (i)
the selection of habitat variables to measure, (ii) the
accuracy (or error) of measurements, and (iii) the procedure of correlation of population and habitat attributes.

(i) Selection of habitat variables

If we accept that the habitat of an organism is its physical
and chemical environment, in all its complexity, this
results in a potentially immense array of variables which
may affect the abundance and dynamics of the species
under consideration. Typically, the vast majority of these
variables are rejected a priori because (i) they are not
thought to play a role (note that this is a subjective
decision, not tested and usually not based on empirical
evidence), or (ii) they are difficult to measure. Those
variables which are presumed to have the greatest bearing
on the population are assessed, and it is hoped that the
limiting variable(s) are included in this subset. Ultimately, however, we can never know whether our analysis
includes those variables affecting the population or
whether we have truly identified limiting variables,
because to do so would require rejecting all others (i.e.
falsifying the role of all the others), which is not practical
given the large number of variables. This point of
falsification is necessary if we are to call ecology a
science, as the current philosophy of scientific practice is
one of Popperian falsification. Critics to my argument
point out that our methods are sufficiently powerful and
that we do not need to employ rigorous falsification
procedures. This may be so, but then is it still science? If
we do not use falsification, but rather verification, we
must instead rely on intuition and inference that our
selection of variables is of importance to the population,
and this approach has two flaws which are harmful. First,
it presumes that the relationship between an organism
and its habitat is direct and observable. We only select
those variables which, from our human perspective, are
likely to have a direct effect on the organism. We have no
reliable method of measuring/evaluating the possibly
influential indirect effects of habitat (e.g. affecting food
supply or predation rate), or synergistic interactions of
two or more seemingly less influential variables. We can
only select those observed relationships which are
apparent to our human eyes, not necessarily the relationships as experienced by the organism. Second, the use of
prior knowledge to select variables for measurement may
not be useful. It is common to take information from
other geographic areas and apply it to our study area (i.e.
select our variables based on the findings of others), but
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spatially these relationships can, and indeed most

probably do, change and so the limiting variables from
one study may not be limiting, or even applicable, in
another. Thus, when striving to use habitat as more than
simply a distributional descriptor, our ability to select
variables is limited to those that we can see and measure
and these variables may change in importance for a single
species over space (and most likely time as well, requiring
our continual development of modifications of the
habitat concept such as habitat bottlenecks, critical
habitat, etc.). Therefore, our selection of variables for
measurement may, or may not, capture the underlying
relationship between the organism and its habitat.

(ii) Accuracy (error) of measurements

If we are to understand a relationship between organism
abundance/dynamics and habitat (given that one exists)
the measurement of specific population attributes, such
as abundance, growth, birth rate, etc. is required. Any
estimates of these variables will have some associated
error due to our very limited ability to sample populations rigorously. This error is composed of measurement error as well as unknown violations of
assumptions in our sampling protocol (e.g. non-random
or biased sampling) and population estimation models
(e.g. immigration/emigration using a closed population
model). In many (most?) cases, these errors associated
with the estimates will make any fine scale resolution of
abundance with habitat impossible to discriminate.
Population size is not the only variable to be measured
with error. Error will also be associated with sampling
the habitat variables, whatever we select them to be.
However, the habitat measurement error, relative to the
error associated with population size estimates, are likely
to be much lower. Therefore, for the sake of this
argument, habitat measurement error will be considered
negligible though in some situations (e.g. quantifying
spatially highly heterogeneous or temporally highly
variable habitats) it may be significant. The predominant
issue with respect to errors of measuring habitat,
I believe, is the question how do we discretize
the continuous environment that the organism experiences? The physical and chemical environment in
which the organism lives is a continually fluctuating
continuum and so we are forced to impose artificial
distinctions on it in order to allow measurement and
analysis. Weather provides a good example. A terrestrial
organism exists in an environment of daily, seasonal and
lifetime changing conditions of temperature, precipitation, sunshine, etc. To quantify these things we use
human conventions of measure (degrees, cm, mm,
microEinsteins, etc), often on arbitrary temporal scales
(weeks, months). Our measurements are convenient and
amenable to our analysis, but do they truly reflect what
is important to the organism?
OIKOS 110:3 (2005)

(iii) Correlation of population and habitat attributes

Attempts at quantifying organism abundance/density
and habitat attributes are based, generally, on correlation; that is, a greater abundance or density implies more
favorable habitat. However, as Van Horne (1983) points
out, the use of simple presence or abundance may be
misleading as it is possible for large densities of
organisms to exist, at the time of sampling, in poor
habitats (i.e. population sinks). She suggests that rather
than presence, a function of density, survival, and
fecundity be used as a measure of habitat quality,
though this still leaves us with simple correlation
between population attributes and habitat !/ it does
nothing to present mechanistic relationships. We
must keep in mind, this point particularly lacking in
the ecological literature, that correlations may be spurious or misleading !/ they do not indicate causative
relationships between variables. And so, if we are to
rigorously understand organism-habitat relations, correlation is a valuable guide to suggest possible habitat
variables of importance but it remains impossible to
state a causative relationship exists between the two
variables until we can show mechanistic (i.e. biological,
ecological) links.
A related issue, unfortunately much neglected, is that
of time lags. The population that we see/measure at
present is likely to be a function of a previous and
unmeasured habitat condition, not a response to present
habitat measurements. As no population can respond
instantaneously to changes in the environment, the
population we see and measure at the present time is
actually a function of the habitat at a previous time, not
the present one we are sampling. Therefore, we may be
trying to correlate population attributes with variables
which are independent of the population (i.e. did not
play a significant role in affecting the population). In a
similar vein, stochastic events (e.g. mid-winter floods in
streams, unusually extreme cold during a short period of
winter) may go unmeasured in an area and yet have
tremendous effect on a population. But our lack of
knowledge of these events (due to not monitoring) blinds
us to including these most significant events structuring
the populations in our analysis. Either of these factors,
time lags or stochastic events, will not be included in our
standard approach of correlation of current habitat
conditions and populations.
Finally, a correlation of organism abundance/
dynamics with habitat can only exist if the population
is near maximal abundance for that habitat (i.e. near
carrying capacity). If the population is not maximal,
and is indeed varying about a lower value, it will
be controlled by other factors (e.g. predation, disease,
etc.) and so habitat cannot show a relationship
with it. This will show up as the two variables
being uncorrelated. Therefore, correlation only applies
to populations near maximal size. This point immediOIKOS 110:3 (2005)

ately excludes quantitative habitat !/abundance relationships, via correlation, for rare and endangered species
which are, by definition, far below their maximal
abundance.
Therefore, based on the foregoing, there exist fundamental problems regarding our ability to quantify and
relate habitat attributes and population abundance of
organisms. These problems are: (1) our selection of those
variables to be measured is based on inference and
convenience rather than relevance; (2) errors in measurement of variables prevent fine scale resolution of
organism abundance, thereby precluding observation of
changes in abundance with habitat, and (3) the correlation of abundance with habitat does not account for
mechanistic relationships, time lags or stochastic, but
highly influential, events, and is only applicable when
populations are near maximal abundance. These problems have created such difficulties that is has recently
been suggested (Van Horne 2002) that one of the
fundamental components of modeling !/ that of validation using an independent data set !/ be de-emphasized
in habitat modeling. She maintained [habitat] models
cannot be universally validated because they are too
general to make precise and testable predictions, apply
only within tight and often unknown boundary conditions, or have failed to incorporate process and therefore
rely on correlations that may be spurious (p. 65). I
perceive this philosophy as further support of my
contention, as my interpretation of it is that we cannot
develop general models of habitat !/abundance. Thus, it is
conceivable that this is due to such relationships simply
not existing. The following case study, while dated,
further emphasizes this.

A case study
Habitat !/population analysis and modelling have a long
history in freshwater fisheries management, the first
ones go back over 50 years. Therefore, if success in
understanding habitat quantitatively, in terms of effect
on populations, is to be found anywhere, this is likely to
be one of the fields worth examining. The primary
emphasis with regard to habitat in freshwater fisheries
research has been to correlate the standing stock
(numbers or biomass at a given time) and habitat
variables which might then allow predictive estimates
of future, or geographically separated, populations based
on environmental characteristics. In 1988, Fausch et al.
reviewed 99 models constructed between 1950 and 1985.
They found a large number of technical problems with
nearly all of the models, and that those few models (n "/
10) which had attempted validation with independent
data sets were found to have very low predictive ability.
This collection of models involved 30#/ species of
freshwater fish and over 100 habitat variables (measured
and derived), ranging in scale from the microhabitat to
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the drainage basin. Yet despite this impressive amount of

effort there still does not exist clear, unambiguous
relationships of habitat and abundance for a single
freshwater fish species which may be applied to conditions beyond the specific location and time of sampling.
Fausch et al. list a series of technical problems with data
collection and modelling, but I propose that their
findings are evidence of a more fundamental problem
!/ that habitat, as we choose to define and measure it, is
not related in a quantitative way to species abundance.

Implications
If the thesis developed here is correct, this re-evaluation of habitat would have significant effects on all
aspects of our understanding of basic and applied
ecology, and the management of species. It implies that
(at a minimum)
1)

2)

3)

Understanding population dynamics and ecology of

a species without a fuller approach, including biotic
factors such as competition and predation, is not
possible. Trying to understand or predict abundance
of organisms solely by correlating environmental
conditions will fail. Interestingly, back in 1959,
Udvardy thought about habitat as an abstraction
of the essential physical factors and of the essential
co-inhabitant biota, in a locality where individuals
of that population regularly live and reproduce
(p. 725; emphasis mine). I believe that he was on the
correct course and we have been waylaid when we
neglected the biotic aspect of this concept.
Habitat modelling of a species is of value only
under the special case where habitat is limiting the
populations and this is likely to be spatially and
temporally bounded. In the general case, the
modelling exercise may provide insight into allowable space for a population to expand into, given
the relaxation of mortality factors, but cannot
realistically depict the present or future state of
the population under naturally occurring conditions.
The single-minded emphasis on habitat protection
by management agencies over the last 30 years has
distracted us from utilizing other approaches which
may provide greater information on factors affecting abundance and so developing effective protec-

tion and recovery strategies. Based on the preceding

pages, I would argue that the provision or protection of habitat alone will not influence populations
if that is not the factor which is controlling them.

Final word
It has been my intention to try to build a strong
argument against the habitat concept in order to provoke
debate; I have perhaps overstated some points as a form
of further provocation. I believe the only way we will
move ahead is to constantly challenge our underlying
assumptions, and each other. My goal is to foster some
deeper thinking on this fundamental tenet of our science
and I hope I am successful in encouraging others to
critically evaluate this cornerstone of our beliefs.
Acknowledgements !/ I would like to thank Barry Taylor, Doug
Rowland, Doug Peterson and Raimo Virkkala for thoughtful
comments/criticisms on a draft of this essay. Their perspectives
on this provided some insights that I had not considered.

References
Fausch, K. D., Hawkes, C. L. and Parsons, M. G. 1988. Models
that predict standing crops of stream fish from habitat
variables: 1950 !/85. !/ USDA, Pacific Northwest Research
Station, General Technical Report PNW-GTR-213.
Hall, L. S., Krausman, P. R. and Morrison, M. L. 1997.
The habitat concept and a plea for standard terminology.
!/ Wildl. Soc. Bull. 25: 173 !/182.
Looijen, R. C. 1995. On the distinction between habitat and
niche, and some implications for species differentiation.
!/ Poz. St. Phil. Sci. Human. 45: 87 !/108.
Morrison, M. L. and Hall, L. S. 2002. Standard terminology:
toward a common language to advance ecological understanding and application. !/ In: Scott, J. M., Heglund, P. J.,
Morrison, M. L. et al. (eds), Predicting species occurrences.
Issues of accuracy and scale. Island Press, pp. 43 !/52.
Odum, E. P. 1971. Fundamentals of ecology, 3rd ed. !/ W.B.
Saunders Company.
Udvardy, M. F. D. 1959. Notes on the ecological concepts of
habitat, biotope and niche. !/ Ecology 40: 725 !/728.
Van Horne, B. 1983. Density as a misleading indicator of
habitat quality. !/ J. Wildl. Manage. 47: 893 !/901.
Van Horne, B. 2002. Approaches to habitat modeling: the
tensions between pattern and process and between
specificity and generality. !/ In: Scott, J. M., Heglund,
P. J., Morrison, M. L. et al. (eds), Predicting species
occurrences. Issues of accuracy and scale. Island Press, pp.
63 !/72.
Whittaker, R. H., Levin, S. A. and Root, R. B. 1973. Niche,
habitat and ecotope. !/ Am. Nat. 107: 321 !/338.

Subject Editor: Jan Lindstro

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