Chaos, Solitons and Fractals 12 (2001) 197203

www.elsevier.nl/locate/chaos

Preface

Chaos in ecology
Vikas Rai a, W.M. Schaer b
a

Structural Biology Unit and Computer Centre, National Institute of Immunology, Aruna Asafali Road, New Delhi 110 067, India
Department of Ecology and Evolutionary Biology, Program in Applied Mathematics, University of Arizona, Tucson, AZ 85721, USA

Overview
As noted by Hao Bai-Lin in the preface to his admirable collection [29,30] of inuential papers on nonlinear dynamics, the discovery [48,55,56] of chaos in ecological dierence equations, as much as anything
else, fertilized a owering of interest in this subject some twenty-ve years ago. Perhaps not surprisingly, it
was in the physical, as opposed to the biological, sciences that ``chaos theory'', as it is often (and inaccurately!) referred to, really took hold. In ecology itself, the ubiquity of chaos and other non-linear phenomena in both discrete and continuous models was subsequently conrmed 1 [1,3,4,9,27,34,46,57,7072].
At the same time, convincing evidence for chaos in natural systems proved harder to come by [17,33,64].
For example, in the case of pre-vaccination epidemics of measles in large, rst world cities, what was once
judged [63] to be one of the more likely examples of real-world ecological chaos, is now the subject of
divergent opinion [16,18].
In as much as ecological systems have all the necessary ingredients: palpable non-linearity, multiple
state variables, etc., for chaotic dynamics, the lack of evidence for chaos in nature may strike some as
surprising. The conventional view [64] is that ecological data sets are too short and too corrupted by
observational error and process noise to allow for accurate characterization of the underlying dynamics.
This has prompted a search for better methods of extracting the deterministic signal from noisy time
series [17,19,20,58,8082]. The inherent diculty of this task is underscored by two facts: in the rst
place, as emphasized by Auerbach and others [7,47], chaotic motion can be viewed as a choreography
involving non-stable cycles, with the lead dancer, the evolving orbit, successively favoring dierent
partners. It follows that chaotic time series will often have strong periodic components rendering it
dicult to distinguish them from null models [17] consisting of periodic motions in the presence of
noise. 2
An additional problem results from the fact that the time evolution of non-linear systems is often
reective not only of the attractors to which trajectories tend in the limit of large time, but also of the
presence of non-stable sets elsewhere in the phase space. 3 For example, in the case of the Lorenz
equations, Yorke and Yorke [87] have pointed out some time ago that there exist parameter values for

Many of the early observations [2,6,37,75] of complex dynamics in ecological models were published in physics and applied
mathematics journals where they remain, to this day, unknown to most ecologists.
2
Such diculties are exacerbated by what has been called [40,41] ``transient periodicity''. In this case, a chaotic attractor contains
well-dened semi-attractors on which the motion is semi-periodic with the consequence that episodes of ``noisy periodicity'' [51] are
interspersed with periods in which the dynamics are more obviously aperiodic. In the case of short time series in which there occur only
a few such transitions, the naive observer is likely to suspect some undetected change in environmental conditions as causing the
change in behavior.
3
The reality of non-stable invariant sets, and their inuence on observable dynamics, have recently been conrmed in the case of
our beetle dynamics in the laboratory [13].
0960-0779/00/$ - see front matter 2000 Published by Elsevier Science Ltd. All rights reserved.
PII: S 0 9 6 0 - 0 7 7 9 ( 0 0 ) 0 0 2 1 3 - 7

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which one can observe arbitrarily long chaotic transients even though the dynamics tend asymptotically
to equilibrium. This phenomenon, which they termed ``metastable chaos'', results from the fact that
coexisting with a pair of stable xed points is a chaotic saddle (Sparrow [78] calls it a ``strange invariant
set''.) on which the motion is every bit as chaotic as on a bona de strange attractor. When such systems
are subjected to random shocks without the parameter values, there arises the possibility of what Kantz
and Grassberger [39] call the ``noise-stabilized chaos'', i.e., the system spends most of its time in the
vicinity of the chaotic saddle even though the deterministic behavior is simple. 4 In such cases, eorts to
ferret out the asymptotic dynamics miss the essential point i.e., that it is the saddles, not the attractors,
that determine the motion.
An alternative point of view is that chaos in nature really isa rarity and that this reects something
fundamental about the organization of ecological systems. One possibility [65,83] is that the reality of
ecological interactions, their non-linear character notwithstanding, necessitates high levels of dissipation which, in turn, preclude the possibility of chaotic motion in the absence of noise. Another [10] is
that chaotically uctuating populations are more prone to extinction than equilibrial populations with
the consequence that group selection acts to eliminate species which would otherwise evolve into
chaos. 5
Contributing to the laggardly rate of progress in non-linear ecology is the fact that natural systems are
not readily brought into the laboratory. An exception is the our beetle, Tribolium sp., an agricultural pest
that has infested mankind's granaries since the days of the ancient Egyptians [77] at least. In this case, the
laboratory system (one or more species in a jar containing our) is not unlike the natural one (one or more
species in a silo). Moreover, the simplicity of the system, and the fact that it is well understood, allows for
the formulation of tractable models whereby quantitative predictions can be tested under a variety of
experimental conditions. Gratifyingly, many of these predictions, including the observation of chaos, have
been borne out by the experiments [1315]. In short, the our beetle system allows ecologists to do what
workers in other elds often take for granted: to write down mechanistic models which predict sequences of
dynamical behaviors that are obtained in response to variations in parameters and then to determine
whether or not the said sequences are actually observed. The results of the beetle experiments, which are in
the time-honored tradition of Gause [25], Nicholson [61], Huaker [36], etc., cannot, of course, be applied
directly to the eld. What they can do, however, is to give plausibility to the idea of modeling ecological
systems mathematically, a proposition which is not well accepted by ecologists and eld biologists.
Moreover, the experiments establish in an undeniable way the reality of non-linear phenomena in the
context of population biology.
The prevailing uncertainty regarding complex dynamics in nature has hardly dampened the enthusiasm
of theoreticians intent on elucidating the possible consequences of non-linearity to the natural world. In this
vein, Allen and his associates [5,67] have challenged the assertion that group selection acts to eliminate
species which uctuate chaotically. Drawing on the observation that most species consist of semi-isolated
populations, these authors argued that exponential divergence of nearby trajectories will often reduce the
likelihood of ensemble, i.e., of species, extinction that would otherwise result from the vagaries of a uctuating environment. From the perspective of species persistence, chaos in nature, by this reckoning, is
good.
The protective eect of chaos observed by Allen et al. is a consequence of sensitivity to initial conditions
[69,76], the dening characteristic of chaotic motion which, in this case, acts to counter the eect of migration which would otherwise act to synchronize spatially disjunct populations. However, as pointed out
by Farmer and his associates [21], there are dierent levels of sensitive dependence. In the case of the

Closely related are the supertransients which occur in spatially extensive systems [32,38]. In this case, one also observes extended
episodes of chaotic behavior before the system nally settles down to its asymptotic state typically a xed point. From a
phenomenological perspective, supertransients are distinguishable from transient chaos in spatially homogeneous systems by virtue of
the fact that transient chaos appears to obtain over open intervals of parameter values.
5
The argument depends on the observation that in simple models, such as the logistic and Ricker [66] maps, minimum population
sizes decline as one enters, and then moves farther into, the chaotic region. For an earlier, ``pre-chaos'' example of such reasoning, see
Gilpin's [26] monograph on the evolution of predatorprey systems.

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199

R
ossler band [68], for example, time series for two of the three variables are characterized by power spectra
in which ``instrumentally sharp'' peaks are superposed on a noisy background. 6; 7 Farmer et al. have
termed such a motion as ``phase coherent'', and it is not surprising that coupling two or more phase coherent systems spatially leads to correlated dynamics in space. Indeed, just such a ``phase synchronization''
has been reported by Blasius et al. [11,50]. In this study, a three-level food chain model with R
ossler-like
dynamics is assumed to determine the within-patch behavior, and local populations are coupled by migration. Of ecological interest is the fact that the system's behavior is reminiscent of the lynx-hare cycle [23]
in that variations in cycle amplitude exceed variations in period. Also of interest is the authors' suggestion
that the high degree of synchrony among lynx populations across North America is reective of phase
coherent chaos 8 in local populations.
Wildlife's ten-year cycle [42] in Canada's north woods is the subject of two other recent investigations.
Reviewing lynx fur returns for the Hudsons Bay Company, Gamarra and Sole [24] have argued in favor of
an earlier suggestion [70] that the system underwent a bifurcation, possibly in response to increased
trapping pressure, in the early 19th century. With regard to contemporary cycle dynamics, a realistic (as
these things go) model of snowshoe hare demography was recently proposed [44] which induces 812-year
oscillations for parameter values that fall within the ranges of independent estimates deriving from eld
observations. 9 Of course, the eld-derived estimates are crude, often being no better than the order of
magnitude, with the consequence that the observed concordance of theory and observation is hardly denitive. Nonetheless, this eort marks the rst successful attempt to model what is arguably ecology's mostcelebrated oscillation in a falsiable manner.
Of more general import is the fact that the snowshoe hare model can be formulated as a perturbation of
a Hamiltonian limit wherein arise the motions of ecological interest. The proposed restoration [43] of
Hamiltonian dynamics [35,49] to a central place in ecological theory is both novel and very much against
the conventional wisdom [54] which holds that conservative systems [84,85], by virtue of their structural
instability, are irrelevant to the behavior of realistic models. The aw in this point of view turns out to be
the structural stability criterion itself, which is based on the notion of topological conjugacy. Essentially,
many of the orbits of structurally unstable systems in the present case, an innite number can persist
under perturbation over open intervals of parameter values. The surviving motions include periodic orbits
that give rise to stable cycles in the dissipative regime and probably also to the saddle cycles about which
chaotic attractors are organized [43].
About this volume
The foregoing (and other) developments, the historical importance of ecological theory to non-linear
dynamics, and the authors' personal opinion that non-linearity will eventually be accepted as a fundamental property of ecological systems, motivated the decision to produce a special issue of Chaos, Solitons
and Fractals on chaos in ecology. Each of the papers included addresses an important topic. We now devote
a few sentences to each.

By ``instrumentally sharp'', we refer to peaks as narrow as the sample size permits. As in the case of R
ossler's equations, the degree
of phase coherence in tri-trophic level ecological models [34] is parameter-dependent. Recently [43], it has been pointed out that the
origins of this dependence lie in the fact that such systems can be formulated as perturbations of one or more Hamiltonian limits [35,49]
from which emanate resonance horns corresponding to cycles of all possible base rotation numbers. With increasing levels of
dissipation, the cycles are destroyed by saddle node bifurcations in a well-dened sequence. Phase coherent chaos results when the
cycles, about which a chaotic attractor is organized, are all associated with a single resonance horn. Since all the cycles have the same
base period, it follows that the power spectrum will have sharp peaks and that, from a time series point of view, variations in cycle
amplitude will exceed variations in period.
7
Animations illustrating the mixing properties of the R
ossler band and other chaotic attractors may be viewed and downloaded at
http:bill.srnr.arizona.edu. (Click on Cool Chaos Demos.)
8
An alternative explanation, of course, is periodicity in the presence of noise.
9
Information on the ``ten-year'' cycle generally and the snowshoe hare model, in particular, is available on-line at http://
two.ucdavis.edu/aking/mam99.

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The paper by Rai and Upadhyay attempts to understand the general failure in the eorts to observe
chaos in natural populations. For two representative models, it is shown that the range of parameter values
corresponding to chaos is both narrow and biologically unrealistic. It is further observed that for those
parameter regimes in which chaos does obtain, the generative mechanism involves catastrophic bifurcations
known as ``crises'' [28].
The contribution of Cushing et al. summarizes the aforementioned studies of laboratory populations of
the our beetle, Tribolium castaneum. The methodology is both elegant and rigorous. In the rst place, a
deterministic model is shown capable of inducing time-series which qualitatively match those obtained
experimentally. Quantitative correspondence is then achieved by replacing the deterministic model with its
stochastic counterpart thereby providing for the eects of demographic and environmental noise. As noted
above, this work has yielded the strongest evidence to date for chaos in an ecological system. Together with
the interdisciplinary approach (modeling, experimentation and statistical analysis), it will likely serve as a
model for future investigations of ecological population dynamics.
The third paper by Gamarra et al. focuses on chaos control [22] and techniques which can be used to
identify chaos in non-linear population models. The authors demonstrate that their methods are robust to
environmental stochasticity and can further be applied to individually oriented studies. Of the various
chaos control strategies currently available, Gamarra et al. identify the one best suited for controlling chaos
in realistic models. Further research along these lines may also facilitate the detection of chaos in natural
data.
The next two contributions focus on periodically forced predatorprey interactions. In the rst, Schaer
and co-workers argue that the analysis of Hamiltonian systems can help explain cyclic phenomena in nature
(see above). In particular, they review the mechanism by which periodic motions arise in the Hamiltonian
limit and point out that empirical observations [12] of allometric scaling (cycle period versus body size) in
mammals result as a natural consequence. The second paper by Vandermeer and colleagues uses circle
maps to describe the dynamics of a periodically forced LotkaVolterra system. As predicted by earlier
investigations [43,45], coexisting attractors are observed. Vandermeer and his associates further observe
that the system is characterized by fractal basin boundaries [52,53] and that, in some instances, the basins of
attraction additionally manifest the so-called ``Wada'' property [62]. These matters are further discussed
with regard to the model (see above) studied by Blasius et al. [11].
The next two papers concern aquatic systems. The study by Tikhonov et al. discusses the dynamics of
schooling behavior in sh using reaction-diusion equations. Two types of behaviors are distinguished:
motions of low-persistence and behaviors which are highly persistent. In the former instance, fractals on all
time scales are obtained. In the latter, multi-fractals are observed for large-scale displacements. The
movement patterns of sh are of obvious consequence to the organisms they eat which, depending on
the size of the sh in question, are either smaller sh or zooplankton. These consequences are the subject of
the paper by Edwards and Bees. The latter authors consider the dynamics of tri-trophic plankton models 10
which utilize ``closure terms'', [79] of the form dZ m , to model the eects of sh predation on the zooplankton concentration, Z. This device allows for the omission of sh population density as an explicit
variable. Historically, linear closure terms have been used to model ``passive'' predators, for example, lter
feeders, and quadratic terms, to model predators which are attracted to the regions of high prey concentration. Arguing that the fractions of predators employing alternative feeding strategies vary with environmental conditions such as turbulence, Edwards and Bees consider non-integer values of the ``index of
closure'', m. Among other things, they nd that dynamical complexity observed for the special case, m 2,
is persistent over a range of values in the interval, m 1; 2.
Ecological exploitation is also the subject of the paper by Bernard Cazelles who establishes that it is
possible to construct simple predatorprey models with multiple attractors and riddled basin boundaries
[86]. The practical signicance of this special type of boundary is that even qualitative predictions regarding
a system's time evolution are impossible.

10

The state variables are concentrations of nutrients, phytoplankton and zooplankton.

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The paper by Rinaldi et al. addresses the problem of forecasting successive maxima in ecological time
series. As foresters and their entomological colleagues will readily attest, this is a matter of practical, as well
as theoretical importance. 11 The authors focus on the next amplitude maps which, when the dynamics are
on a two-torus or a low-dimensional strange attractor, aord a straightforward device for predicting the
timing and amplitude of the next irruption. The utility of this scheme, which has the practical advantage of
requiring limited information about the previous outbreaks, depends on the prevalence of low dimensional
ecological motion. In essence, this work takes us back to earlier attempts [73,74] to use such maps to
distinguish chaos from random uctuations.
Not unconditionally, the problem of distinguishing deterministic from stochastic dynamics is the subject
of the paper that follows. In this contribution, Kendall considers data generated by a discrete hostparasitoid model in the presence of noise. Kendall compares the ability of a non-parametric model to
describe the data with that of the original equations. He concludes that the original model outperforms the
non-parametric scheme which, in turn, outperforms a second incorrect model chosen for comparison.
Kendall also observes that oscillatory transients provide sucient structure for parameter estimation and
identication of the correct model and that the non-parametric model is most readily rejected when the
dynamics are chaotic. This analysis, which is in the tradition of Morris [59] and other early investigators
[73], by virtue of its situation-specic nature, underscores the diculties which attempt to formulate a
general methodology for detecting complex determinism in the absence of the ability to conduct controlled
experiments.
The nal paper is by Jon Allen and his associates. The model considered here belongs to a distinct class
known as ``discrete spatial convolution'' models, whereby the eects of migration and dynamics at a point
are studied in the context of spatially extensive systems. Specically, the authors conrm the stabilizing
eects of spatial structure for a hostparasitoid system under various assumptions. In addition, the analysis
of spatial convolution models via the frequency response of the system transfer function is illustrated. Realworld systems are, of course, distributed over the surface of a two- (and sometimes three-) dimensional
planet, with the consequence that the ``well-stirred'' dynamics of discrete maps and ordinary dierential
equations are always an approximation. Often, it is also a poor one for example, when the spatially
extensive generalization of a non-spatial model is characterized by spatial structures [31] or supertransients
[32,38]. It is therefore tting that the concluding contribution to this volume takes up the problem of
ecological dynamics in space, as well as time.
Acknowledgements
We thank Tatiana Bronnikogva, Bob Costantino and Aaron King for their criticism and comments. The
mistakes, of course, remain our own.
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We refer to the irruptive dynamics of insect defoliators, spruce budworm, larch budmoth, etc., of north-temperate forests. These
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