Oecologia (1991) 86:144-145

Oecologia

9 Springer-Verlag 1991

The usefulness of the equilibrium concept in population dynamics

A reply to Berryman
Henk Wolda
Smithsonian Tropical Research Institute, APO Miami, FL 34002~0011, USA
Received June 13, 1990 / Acceptedin revised form October 30, 1990

I am grateful to Berryman (1990) for trying to help with

my dilemma (Wolda 1989), but I am afraid that his note
does not solve my problem about the practical meaning
of the equilibrium concept. I see no way to measure
an equilibrium in the field, as it seems impossible to
separate fluctuating equilibrium values from fluctuating
deviations from those equilibrium values. Population
density at a given moment may be high or low relative
to densities at other times, but there is no way of knowing whether that density is higher or lower than the equilibrium value at that time. This questions the validity
of density dependence tests that rely on high densities
being above and low densities being below equilibrium.
Berryman (1990) does not refute any of this, but he
does make some interesting observations that require
an answer.
Berryman claims that density dependent regulation
can occur without the existence of a stable equilibrium
(a point attractor) and that the idea that stable equilibria
are a necessary condition for density dependence to exist
is illogical, disagreeing with much of the literature I
know. I do not understand his argument, but, I fail to
see how density dependent regulation can be anything
but a process that tends to reduce population size when
high, higher than the present equilibrium value, or range
of values, and to increase population size when it is
lower than the equilibrium at that time. Berryman gives
some examples of equilibria emerging as a result of an
analysis. However, if I read the literature right, the test
he uses is invalid (Maelzer 1970; St. Amant 1970; Reddingius 1971; It(5 1972; Pielou 1974) and there seem to
be problems with all other existing tests (Solow 1990;
Reddingius 1990; Den Boer 1990b). Moreover, the fact
that for each population Berryman finds one stable,
time-independent, equilibrium point makes the analysis
suspect. How can an equilibrium in a natural population, set by a complex of varying environmental processes, fail to vary in time? How, if at all, does a constant
equilibrium point, found when applying some model to
population data, relate to a more realistic time-varying
equilibrium in that population?

Berryman misses my point when he states that I conclude that an equilibrium has to vary because population
size fluctuates. I claim it has to vary because its value
is determined by the environment and its interaction
with the organism. Most pertinent environmental factors
in natural habitats vary, often a great deal, which is
why I do not see how an equilibrium, assuming it exists,
can be constant. Berryman's discussion of various possible causes of population fluctuations other than a varying equilibrium thus seems irrelevant. However, Berryman is right when he states that the fact that equilibrium
structures can be influenced by time-varying factors
makes their estimation difficult does not invalidate the
concept of density dependence. I never claimed it did.
I never even claimed that density dependent regulation
does not operate in natural populations. It is only that
I am not willing to accept that it does without proof,
and such proof seems difficult, if not impossible, to obtain. I do not agree that the difficulty, or even impossibility, of estimating fluctuating equilibrium values is not
an ecological problem, even if methods used are statistical.
Berryman does not recognize the difference between
"stabilization" and "regulation" as defined by Reddingius and Den Boer (1989). Their definition is: "The
often supposed (general) tendency for population density to stay for some time between relatively narrow limits will be called 'stabilization' [..]. Stabilization may,
but need not, result from the influence of'governing'
(density dependent) factors [..]. Only if this is the case
it shall be called 'regulation'." This because he cannot
see stabilization to occur without regulation. With Royama (1977) he accepts density dependent regulation as
a basic truth logically derived from the persistence of
natural populations. However, I and many others have
observed populations in apparently stable natural surroundings change dramatically in abundance. Populations of some originally common species dwindled to
extinction while others, originally rare or even absent,
became common, which hardly suggests regulation or
even stabilization. The "persistence" of natural popula-

145
tions is not necessarily a fact, which means that neither
is regulation. On the other hand, those of us who no
not strictly believe in density dependent regulation as
a "basic truth", an axiom, a dogma that does not need
to be proven, recognize the possibility that completely
unregulated populations not only may exist but may
stay within limits over a considerable period of time.
That is, such populations do not necessarily reach catastrophically high densities, or become extinct, while we
watch (cf. Reddingius 1971). Berryman's contention that
stabilization and regulation have the same underlying
causes is not necessarily correct and I maintain that it
is conceptually profitable to continue to distinguish between 'stabilization' and 'regulation'. When stabilization is observed, one should keep an open mind and
investigate it causes rather than jump to conclusions
about it being p r o o f of regulation. Many ecologists keep
trying to find a way out of this quagmire of a seemingly
untestable hypothesis that is so basic to the theory of
population dynamics by continued data analysis and review of available evidence (e.g. Strong 1984; Stiling
1988; Hassell et al. 1989; Latto and Bernstein 1990; Den
Boer 1990 a, b).
Berryman's theoretical arguments and references to
negative feedback mechanisms in other systems fail to
convince me. "Negative feedback loops are innate to
all ecological systems" he says, as exemplified by " i f
organisms feed on each other negative feedback loops
occur automatically at the population level." I fail to
see the relevance of A eating B for the question of regulation of numbers in populations of A or B. There is ample
evidence that density dependent processes operate in
natural populations, but this is not sufficient evidence
for regulation to occur (Reddingius 1971; Reddingius
and Den Boer 1989). Berryman blames the current confusion about regulation etc. on " o u r refusal to recognize
that ecosystems obey the same rules as all other dynamic
systems, both natural and engineered." It would simplify
things considerably if they did, but do they? What factual evidence supports this contention ? Non-linear dynamics and general systems theory are undoubtedly useful
theoretical tools, but do they solve the problems at hand
of the practical ecologist?
There is, unfortunately, a wide gap in ecology between theory and practice. Berryman heavily relies on
theoretical constructs for his arguments, while I am more
pragmatic and insist in practical evidence before I accept
theoretical results. Admittedly, theory moves briskly
ahead, but whether much of this theory is relevant for
natural populations remains to be seen. I find the hy-

pothesis of density dependence regulation very attractive

and would like there to be evidence conducive to its
acceptance, but thus far such evidence remains elusive.
This does not mean that population dynamics is at a
standstill. I agree with Berryman that the analysis of
population data does not need the equilibrium concept.
There are many profitable ways to collect and analyze
interesting information on the dynamics of populations
without mentioning the words equilibrium or, for that
matter, density dependence.

References

Berryman AA (1990) Stabilization or regulation: What it all means.

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