Ecological Modelling 117 (1999) 305 – 314

Mastering chaos in ecology

Inti Suárez *
Zoologisches Institut der Uni6ersität Basel, Rheinsprung 9, CH-4051 Basel, Switzerland

Received 3 November 1997; accepted 15 December 1998

Abstract

Recent developments in dynamical system theory consider chaotic fluctuations of a dynamical system as highly
desirable behaviour because fluctuations allow such a system to be easily controlled. This recent development, chaos
control theory, has been tested in several physical situations. The intrinsic instability of the orbits embedded in a
strange attractor make them easy to change to a more stable and predictable behaviour. The existence of chaos in
natural populations, at least in some species of pest insects, makes the topic of importance in pest management. The
techniques proposed to achieve control over a chaotic dynamical system are presented and discussed in an ecological
context. An important point in the application of this technique to real systems is the amount of noise involved in
the estimation of the parameters of the system. It is also important in the determination of the perturbations required
in the control parameters of the system. The theoretical possibility of obtaining these estimations in field population
biology provides opportunities for using these techniques in pest management. The control of chaos theory shows
how chaotic systems can be controlled by stable and linked systems. The possibility that chaotic behaviour has been
seldom found in nature because of the existence of linked phenomena is discussed. © 1999 Elsevier Science B.V. All
rights reserved.

Keywords: Chaos control; Chaos detection; Pest control; Complex population dynamics

1. Introduction previously as bewildering behaviour, can be desir-

Amidst the recent advances in the study of
chaos, the algorithm developed by Ott et al.
(1990a) has become one of the seminal papers in
the area. It has been said that chaos, considered
* Tel.: +41-61-2673486; fax: + 41-61-2673457.
E-mail address: suarez@ubaclu.unibas.ch (I. Suárez)
able and advantageous since it has been demon-
strated that a system sensitive to initial conditions
is highly controllable (Ditto and Pecora, 1993).
This idea has been explored in an evolutionary
context by Doebeli (Doebeli, 1993, 1995a,b) and
in medicine by Weis et al. (1993). The object of
this communication is to explore the conse-
quences of these developments in population ecol-
ogy, starting from the uses in other contexts.

0304-3800/99/$ - see front matter © 1999 Elsevier Science B.V. All rights reserved.
PII: S 0 3 0 4 - 3 8 0 0 ( 9 9 ) 0 0 0 0 7 - 1
306 I. Suárez / Ecological Modelling 117 (1999) 305–314
2. Proposed techniques
The central idea of the method proposed by Ott
et al. (1990a) is to stabilize one of the orbits of the
system’s attractor by making a series of small
perturbations on a control parameter. This is
possible because there is an infinite number of
unstable periodic orbits densely embedded within
the attractor. The proposed algorithm, in general,
is as follows:
1. The system has to be studied through Poincarè
sections in order to choose a desirable orbit to
be stabilized.
2. With the reconstruction of the attractor in
phase space, the direction the system will fol-
low in the presence of a determined change in
the control parameter has to be predicted,
using the techniques proposed by Lathrop and
Kostelich (1989).
3. The change in the control parameter that
tends to stabilize the desired orbit has to be
made, monitoring the result in phase space, to
plan the next change.
It has been demonstrated than the fast conver-
gence of this algorithm to the desired orbit is a
consequence of the instability of the orbits in a
chaotic system (Ott et al., 1990a) and this has
been validated in many different experimental sit-
uations (Bayli and Virgin, 1994).
There are some variations in this method, as
follows:
1. Change a dynamic variable from the system,
not a control parameter from the system. (De-
layed controlling feedback, Pyragas, 1992; dis-
sipative feedback control, Rulkov et al., 1994).
This group of techniques presents less practical
complications than the original one.
2. Disturb the control parameter in a periodic
way, e.g. sinusoidal, (Lima and Pettini, 1990),
or with the perturbations in reference to a
determinate value of the parameter (Hunt,
1991).
These particular techniques do not need the
computational resources that the Ott et al.
(1990a) algorithm does, but their validation is less
general. They are more suitable to systems with
high frequency because they do not need a real
time computer analysis of the system, and they
seem to be more robust to noise (Meucci et al.,
1994). They will be discussed later.
3. Chaos control and pest management
Although the techniques of chaos detection
have been improved (Schaffer, 1984; Sugihara and
May, 1990; Ellner and Turchin, 1995), the exis-
tence of chaos in natural populations is still con-
troversial, and several reviews about it have been
written (e.g. Hasting et al., 1993). Following
Pimm (1984) the first attempt to detect chaotic
behaviour in nature was important in shifting the
opinion of ecologists against its existence (Hassell
et al., 1976). Nevertheless Hassell et al. (1976) did
find evidence of chaotic behaviour in a pest insect
population, Zeirapthera fagi (Nicholson’s
blowfly). There is some more evidence that it
could occur in pest populations (e.g. Phyllaphis
fagi, Turchin and Taylor, 1992). Then, consider-
ing that chaos exists in at least some pest insects,
would it be possible to control these populations
through the control techniques mentioned above?
This is the first question that the author intends to
answer.
Thinking about the applicability of the tech-
niques presented above, one that needs a change
in a dynamical variable of the system can be
compared with a planned harvesting of the popu-
lation (M. Doebeli, personal communication). For
example, to change a dynamical variable in the
system could be to change the population size as
convenient (Guemez and Matias, 1993). Because
the use of this technique is studied elsewhere
(Doebeli and Ruxton, 1997) it will be not dis-
cussed here. In an ecological context, the main
advantage of this technique is not needed, because
computer analysis in real time is not a problem,
since the frequency in an ecological system is days
or months, but not nanoseconds. This means that
the human controller can have at least days to
make the numerical analysis and decide the next
perturbation, whether the system goes on and
cycles enough times to allow the next optimal
control. A more relevant point in choosing the
appropriate technique is the sensitivity to the
noise present in the determination of the attrac-
 I. Suárez / Ecological Modelling 117 (1999) 305–314
tor. With regard to this topic Ott et al. (1990b)
point out that in the presence of noise, the ampli-
tude of the perturbation that will be made in the
control parameter must exceed the error length. If
it is possible to keep the noise bound, it was
shown that the typical outbursts of a chaotic
system can be considered as a low probability tail
case (Ott et al., 1990a), occurring rarely. The
technique proposed by Pyragas (1992) is robust to
noise. It uses feedback control. If noise is present
in the system determination, the feedback in-
creases as the noise increases, achieving control in
spite of the noise (Pyragas and Tamasevicious,
1993). Whatever be the choice between Ott et al.’s
(1990a) or Pyragas’s techniques, the possibility of
deciding the amplitudes of the perturbations as a
function of the size of noise needs to be
considered.
To know these amplitudes is, at least theoreti-
cally, possible in population field biology. To
obtain it there are standard techniques to estimate
confidence intervals in the determination of popu-
lation sizes, through capture-recapture methods,
or another standard technique (Krebs, 1989). Al-
though a confidence interval, this is not a direct
measure of the noise involved in population size
determination, but it could be used as an indirect
measure of it.
Reviewing the proposed techniques, resonant
parametric perturbation (Lima and Pettini, 1990)
still has to be considered. The control used would
have a determined amplitude and frequency, but
the problem here is that it seems difficult to make
perturbations with well-determined values artifi-
cially, in an ecological system.
Until now there have been presented here some
criteria to choose an adequate technique, and the
possibility of estimating the noise in ecological
control has been mentioned. The possible exis-
tence of a control parameter that could be manip-
ulated has not been mentioned yet. The logistic
model, a system capable of displaying chaotic
behaviour, has only one control parameter: the
population growth rate. It is obvious that this
parameter is susceptible to manipulation, with
biocontrol agents or insecticides. More flexible
models, such as the Bellows equation, can also be
controlled by perturbing one parameter (Doebeli,
307
1995a). The problem with this kind of manipula-
tion is in determining exactly what is the magni-
tude of the change in the parameter due to the
controller’s perturbation. If for example, the con-
troller applies some amount of insecticide, he will
not know immediately the amount of change in
the rate of population growth., but if he main-
tains a monitoring program, he can estimate this
amount a posteriori, and before making the next
perturbation.
So far, the feasibility of this kind of control in
ecology has been presented. Now, some points
against it will be discussed. Firstly, it must be said
that the small perturbations on the system control
parameters have been permanent in the examples
where the statements discussed above were vali-
dated. This means that once the parameter is
changed from state p to state p’, the latter will not
change to state p’’ or any other until the con-
troller makes the next perturbation. The author
believes that in natural populations it is impossi-
ble to affirm the existence of this steadiness.
Moreover, the opposite cannot be affirmed. The
resilience of natural populations is still a point of
strong debate, with deep implications on manage-
ment ecology and conservation (Pimm, 1984).
Given the disagreements in this debate, the steadi-
ness of the control parameter state will not be
assured until the controlling techniques are ap-
plied to a natural system, and even in that case it
will only validate the statements in that particular
system and cannot be generalized to other natural
systems.
Furthermore, the fast convergence of these
methods has been demonstrated for systems with
steady changes in the control parameter. Thus,
the convergence speed of the control method re-
mains as an open question in ecological systems.
Before passing to a numerical example the pro-
posed algorithms following from the above argu-
ment would be:
1. Obtain historical data on the population to be
controlled. With these, build a time series and
analyze it in phase space, following Schaffer
(1984). It would be useful make additional
analyses to find evidence of chaos, following
Ellner and Turchin (1995). Moreover, such an
analysis permits the derivation of a model of
308 I. Suárez / Ecological Modelling 117 (1999) 305–314
the population dynamics. An estimation of the
parameters of the model to be used is needed,
to identify stable orbits.
2. Make the perturbation in the control variable,
by, for example, the introduction of an insecti-
cide in the insect’s egg-laying locations.
3. Follow the amplitude of the change by stan-
dard techniques of estimation of population
size. Estimate the size of the noise in this
determination to plan the size of the next
perturbation.
4. With the new points in the time series, go back
to the first step.
4. A numerical example
In this section a numerical example of the
control method discussed above is presented. Its
efficiency in controlling a biological population
with unstable dynamics will be shown. In the
present case it will be considered that the determi-
nation of the population density has a stochastic
component, arising from its estimation in the
wild. The control method is the one proposed by
Ott et al. (1990a,b), and it is tried on a simple
model of population dynamics. The model used
was introduced by Maynard-Smith and Slatkin
(1973), and considered by Bellows (1981) to be the
most generally applicable, due to its mathematical
flexibility. The model and the mathematical con-
siderations for the control of the deterministic
case is briefly described, taken from Doebeli
(1995a).
Consider a population with discrete generations
whose dynamics are described by:
Nt + 1 = f(Nt )=Nt
l
(1)
1+(aNt )b
Here Nt + 1 and Nt are population densities in
successive generations. l, a and b are demo-
graphic parameters: l \ 1 is the intrinsic growth
rate, a is a measure of how well the individuals
cope with the environment, and b ] 1 describes
the type of competition that leads to density
dependence (b: 1 could be interpreted as contest,
b\ \1 to scramble competition). The dynamic
of the population density is determined by the
slope of f at the equilibrium N*, given by f(N*)=
N*. The slope is given by the first derivative of
f(N) evaluated at N*, N*\0. As this slope
reaches a critical value chaos sets in. The parame-
ter a is not included in the equation for the slope,
it affects only the equilibrium density, which cor-
responds to the carrying capacity of the popula-
tion, and will be considered as the control
parameter.
Assuming that parameters l and b have values
coding for chaos, it is assumed that it is possible
to manipulate a in each generation, slightly
changing the population ability to cope with the
environment, hence the carrying capacity. In the
case of the control of a pest insect, as discussed
above, these changes could be reduction of avail-
able egg-laying places, or similar perturbations.
Monitoring population size, which has a chaotic
dynamic, a generation in which Nt is close to the
equilibrium value, N* will occur. In this genera-
tion, one can approximate the dynamics of the
system linearly:
(f
Nt + 1 − N*= (N ,a ) · (Nt − N*)
(N t 0
(f
+ (N*,a0) · (at − a0) (2)
(a
Introducing the ability to control the change in
the parameter a by adjusting it linearly according
to the population density
al − a0 = c(Nt − N*) (3)
because c is a constant to be determined, Eq. (3)
could be substituted into Eq. (2), yielding:
Nt + 1 − N*
=
(f (f n
(N*,a0)+ c (N*,a0) · (Nt − N*) (4)
(N (a
If c is chosen to make the modulus of the
expression in brackets less than one, the system
will then approach equilibrium. Evaluating the
) )
partial derivatives, we get the condition
l− 1 b(l−1)(b + 1)/b
1− b −c B1 (5)
l a 20l
The optimal c value occurs when the above
modulus is zero, finally at arriving the equation

I. Suárez / Ecological Modelling 117 (1999) 305–314
l −1 a l 2
0
copt = 1−b · (6)
l b(l − 1)(b + 1)/b
To specify when control has to be applied, we
must provide a criterion to decide when Nt is close
to N*. As can be seen from Eq. (3), the size of the
perturbation depends on this decision. A practical
309
with two major perturbations, much smaller than
the used in Fig. 1a. If it is intended to control a
natural population with this method, an estima-
tion of b and l from the real data is needed, and
a time series of 30 or fewer data points is enough
to provide estimation of these parameters. In the
strategy to decide the value d for which Nt − simulation shown (Fig. 1b) the parameters of Eq.
N*B dcould be to calculate the size of the maxi-
mal perturbation that can be done in each
particular system, accordingly calculating d. Con-
sidering that an uncontrolled chaotic population
dynamic is ergodic, given any d \ 0 eventually a
generation will fulfill the condition (Ott et al.,
1990b).
To include stochastic factors in the simulation,
Nt from Eq. (1) is calculated by adding a random
percentage of itself:
Ntr
Nt = Nt 9 (7)
100
The parameter r is a random number, taken
from a uniform distribution from zero to a maxi-
mal value, fixed at 1, 5, 10 and 20. These extreme
values are reasonable amounts of noise in the
determination of wild population densities.
(Krebs, 1989). The sign of the added expression
was positive with a 0.5 probability each time a
population density was calculated, and negative
for the remaining cases.
Fig. 1 shows the effect of choosing different
values of d, namely 10%, 1% and 0.1% of the
maximal population size. In this case no noise in
the population density is simulated.
Fig. 1a shows the results of the control when
d = 6 (the maximal population density for the b
and l values chosen being 60). In the fifth genera-
tion the control is made active and the equi-
librium density is closely attained after three
relevant perturbations. The equilibrium orbit is
approached asymptotically, so the size of the per-
turbations decreases through time but is not zero.
After the number of perturbations mentioned, the
changes in them occur at the octave decimal
position, and are irrelevant in practice.
In Fig. 1b is shown the result of controlling the
system when d is set to 0.6. After 30 generations
of uncontrolled chaotic dynamics, the condition is
fulfilled and the population density is controlled
(1) can be estimated from the time series before
the control is activated. Nonlinear regression and
the Levenberg–Marquardt parameter estimation
method (provided by SPSS statistical software)
yield estimation of parameters precise to eight
decimal places.
Fig. 1c shows the system with d= 0.06. The
condition to apply the control is only fulfilled
after 1100 generations. It is still noteworthy the
size of the perturbation, which is smaller than in
the preceding cases. Only one relevant perturba-
tion is needed to control the system. In general it
is illustrated that the use of smaller values of d
imply a longer time of monitoring the population
density to start the control, but once it has been
made active, the size of the perturbations and its
numbers are smaller and the technique works
more efficiently.
Fig. 2 shows the control working with different
levels of noise in the population density. The
simulations shown are typical of the system, cho-
sen after many repetitions.
In this case, the technique is robust to signifi-
cant amounts of noise added to the population
density determination. At increased noise levels,
the dynamic is still controlled, but bigger pertur-
bations are needed to keep the system stable.
Even in cases in which the noise is 10% of the
deterministic population density, the method is
still useful for eliminating the fluctuations associ-
ated with a chaotic regime. These levels of noise
are not enough to force the system out of the
neighborhood of a stable orbit. Only in Fig. 2d,
showing a system with 20% noise, is it the case
that the stochastic part of a population’s dynamic
can make the control method no longer useful.
The reason is that such an amount of noise takes
the system out of the neighborhood of equi-
librium. In this case the system returns to chaotic
fluctuations. Out of control again, the condition
to make the perturbation scheme active again
310 I. Suárez / Ecological Modelling 117 (1999) 305–314

Fig. 1. Changing the neighborhood criteria changes the required time for starting the control and the number and size of the
perturbations required. The values of a are shown in the small graphics. The population densities are plotted against generation time
in the main graphics, showing the attainment of the equilibrium density. The parameter values for the figure were as follows:
a0 =0.05, b = 4 and l=5. d= 6 in Fig. 1a, d = 0.6 in Fig. 1b and d= 0.06 in Fig. 1c.
 I. Suárez / Ecological Modelling 117 (1999) 305–314 311

Fig. 2. The importance of noise in the control of an unstable dynamic is considered here. The amount of noise increases from Fig.
2a to Fig. 2d. It is noteworthy that in this noisy case, a constant monitoring of the population is needed to keep its dynamics
controlled. As in the previous figure, the a values are shown in the small boxes and the population densities against generation time
in the main graphics. The parameter values for this figure were as follows: a0 =0.05, b =4, l =5 and d=0.6. r =1 in Fig. 2a, r= 5
in Fig. 2b, r= 10 in Fig. 2c and r = 20 in Fig. 2d.
312 I. Suárez / Ecological Modelling 117 (1999) 305–314
takes some time to be fulfilled, but after it hap-
pens, the system is randomly set out from the
equilibrium again.
This numerical example exemplifies several fea-
tures of the method mentioned above. It is shown
that knowledge about the levels of noise built into
the system to be controlled and the maximal size
of the perturbation to be applied are important
parameters to know before using this control
method properly. It also points to a striking fea-
ture of unstable dynamical behavior. To obtain a
stable orbit from a chaotic system, its parameter
does not need to be changed from chaotic values
to stable values. Small perturbations at the proper
moment are responsible for the change of the
dynamics. For the numerical example shown here
the changes in the control parameter are displayed
in the small windows on the figures. It shows that
the control parameter a is only slightly perturbed.
In the deterministic case a returns to its original
value, but due to the timing of the perturbation
the resulting dynamic is stable and not chaotic.
5. Chaos control and chaos detection in natural
populations:
As mentioned at the beginning, the existence of
chaos in natural populations is a controversial
issue. Without dealing with the methodological
problems for its detection, some arguments
against its existence in nature have been pre-
sented, by means of natural selection (Berryman
and Milstein, 1989). The central argument in this
work is that if a population behaves chaotically, it
will pass through very small population sizes,
increasing the probability of extinction, and there-
fore cannot be favoured by natural selection. Be-
sides, it is clearly a group – selectionist argument
(and makes invalid the point following Ferriere
and Gatto, 1993, among others), and there are
models developed that permit us to consider the
chaotic behaviour of a natural population as an
evolutionary stable strategy, the result of natural
selection acting at the individual level (Ferriere
and Fox, 1995, for a review of their views on the
topic; Doebeli, 1993, and Altemberg, 1991 for
alternative views of chaos as an outcome of natu-
ral selection processes). Allen et al. (1993) argue
that chaotic systems, because of their intrinsic
noise amplification can amplify the noise of the
system, and enhance its natural heterogeneity,
preserving it from extinction.
Forgetting the debate about the existence of
chaos for a moment, let us consider the con-
trolling techniques consisting in the periodic per-
turbation of a main variable of the system (Lima
and Pettini, 1990). Now suppose that the periodic
perturbations occurring in populations, (precisely
in those where chaos was expected to be more
likely, the temperate ones, according to Turchin,
1993) are the periodic perturbations of a variable
of the system. This view can provide a natural
chaos control device in an ecological context, but
there is strong debate about the existence of chaos
in the rodent studied (Turchin, 1993, 1995; Falck
et al., 1995a,b). From this debate, in Turchin’s
original 1993 work are several reasons to expect
chaos in northern populations, but, also in this
northern ecosystem the resources tend to decrease
abruptly during the winter, rising again in the
following seasons, and repeating this pattern in a
periodic way. Perhaps this seasonal pattern of the
environment can be likened to the parametric
periodic perturbation in the method proposed by
Lima and Pettini. Maybe the non-existence of this
behaviour in nature is due to periodic, or seasonal
perturbations, more than the evolutionary argu-
ments against chaos, or the non-existence of
proper detection techniques.
This statement, suggested by one of the chaos
control techniques, agrees with previous works on
metapopulation models. There was controversy
on the existence of chaos in connected metapopu-
lations (McCallum, 1992; Gonzales-Andujar and
Perry, 1993; Ruxton, 1993, 1996a; Doebeli, 1995a,
among others in broader contexts). It hangs on
whether a metapopulation defined as chaotic dis-
plays chaos when connected (through migration)
with another metapopulation whose behaviour
could be, (or not) chaotic. Although there is a
consensus that it depends on the immigration–
emigration levels, these results are reinforced by
the theory developed to explain the synchroniza-
tion and the control of the chaotic systems (Pec-
ora and Carroll, 1990). Furthermore, they are
 I. Suárez / Ecological Modelling 117 (1999) 305–314
becoming established in a broader theoretical
frame, the control chaos theory, as pointed out by
Doebeli (1995b). The fact is that in other disci-
plines it has been demonstrated that two poten-
tially chaotic systems, if coupled, can behave in a
periodic fashion, because of their intrinsic chaotic
natures (Pecora and Carroll, 1990).
As an example of these phenomena in biology,
there are theoretical models of trophic chains that
show chaos (Hastings and Powell, 1991; McCann
and Yodzys, 1994; Eisenberg and Maszle, 1995;
Ruxton, 1996b). A trophic chain implies several
populations connected by predation. If we follow
the suggested idea in the case of a metapopulation
connected by migration we can propose the same
idea. In nature it has become difficult to find
chaotic behaviour probably because the connec-
tions in a trophic chain allow the same regulatory
role for populations with strong periodic be-
haviour. It is important to say that all the authors
cited here discuss to some extent the importance
of coupling as a regulation source, but the point
of this paper is to see this regulation by coupling
enhanced because of the instability of chaos, fol-
lowing Ditto and Pecora (1993).
6. Conclusions
Exploring the Ott et al. (1990a) method and its
variants reveals valuable contributions to the the-
ory of population management. This algorithm is
applicable in the control of pests, insects and
other natural populations, offering a fast converg-
ing solution. The exploration of a simple numeri-
cal example supports this claim.
The existence of linked phenomena in ecology
suggests that the paucity of examples in which
chaos is detected in nature could be due to the
control of chaos by means of periodic fluctuations
from the environment of another population.
Acknowledgements
I would like to thank Luis Miguel Marquez for
comments in the initial version of this paper. To
Michael Doebeli and Dita Vizoso for encourage-
313
ment and continous discussions about the topic.
Two anonymous referees who suggest the addi-
tion of a numerical example, improving the read-
ability of the manuscript.
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