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Resilience and Other Stability Concepts in Ecology: Notes on their Origin,

Validity, and Usefulness

Article · January 2004

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Resilience and other stability concepts
in ecology: notes on their origin, validity
and usefulness1

Per Bodin * & Bo L.B. Wiman

Environmental Science Section, Department of Biology and Environmental Science, Kalmar University, Sweden

Introduction
Resilience – a term that has roots in ancient thinking and was developed in mathematics and
engineering – is slowly coming of age in today’s arena of environmental policy-making.
Attempts to transfer this and related stability concepts beyond their original realms into
domains of ecology and society are far from new. Recent attempts can be looked upon as a
“third wave” (cf., e.g., Lotka 1956/1924, van Dobben & Lowe-McConnell 1975, Holling
1978, Wiman & Holst 1982, NRMK 1983, Wiman 1991, Wiman et al. 1998, WRI 2000,
Folke et al. 2002).

Our scope below is to attempt at establishing a conceptual framework that can help resolve
some basic questions and problems with resilience and other stability concepts in terms of
their validity when subjected to such transfer between realms. Undoubtedly, analogy transfers
between science realms, and between science and policy, can be both powerful and helpful,
and might offer common ground for inter-disciplinary endeavours. However, fundamental
problems will arise if analogy is mistaken for identity. In these notes, we provide a brief
account of the meaning of resilience, and directly related stability concepts, in the realms in
which they were first developed and applied. We then briefly address the transfer of such
concepts that has occurred into the realm of ecology, and some of the conceptual and other
problems that are then encountered. Ways forward towards resolving these problems are
finally indicated, together with a few observations on potentials for conceptual bridges to
“sustainability science” (cf. Lövbrand 2004).

1
Received: 28 May 2004, revised version received: 19 October 2004, accepted: 26 October 2004.
*
Email: [email protected] Tel: +46 480 446252

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Bodin P. & Wiman B. L. B. – Resilience and other stability concepts in ecology

Stability in the mathematics/technology domain

Technical systems almost by definition lend themselves to mathematical modelling tools
that can be applied in various ways, such as in the design of infrastructures (bridges, electronic
communications, energy systems etc.). In this context, it is essential to observe that the term
“resilience” is but one of the several elements that enter into stability analyses. That is,
stability is the overarching concept in the technology domain, not resilience – although, as will
be exemplified later, the latter in certain cases constitute the more important one. Obviously,
as already implied at the outset, any transfer of technology-domain concepts into realms
outside of technology – such as ecological, behavioural or social sciences – would need as
much careful scrutiny as would the transfer of concepts from for instance social science into
the domain of technology or ecology. Such transfers of stability concepts have been advocated
by several researchers (cf., e.g., Lotka 1956/1924, May 1973/1975, Holling 1973, van Dobben
& Lowe-McConnell 1975), but the conditions under which transfers of tools and concepts
can be made have not often been clearly stated. At this point we wish to emphasise one such
condition, namely that systems subject to evolution may not be appropriate targets for
transfers of this kind; this (as will be commented on in more detail later on) is because such
systems can respond to disturbance with structural as well as functional change on the entire
system’s level, so that the model representation itself of the system would undergo (discrete
or continuous) change – taking such structural change into account necessitates a significantly
higher level of modelling as well as of stability concepts than is normally needed for, or even
developed for, technology systems. However, to the extent that evolution can be identified
as, and understood in terms of, mechanisms, forces and principles that belong to an extended
hierarchy of systems – from those with fairly short-term dynamics (ecological time-scales)
to those with long-term dynamics (evolutionary time-scales) and thus including, for instance,
mutation and selection pressures (cf. Haken 1978, Hutson & Vickers 2000) – conceptual
transfers gain validity. But, in brief: not only do systems in nature (and society) change, they
also change how they change, and this property poses a range of challenges to the transfer of
stability concepts between scientific realms.

Also, as an additional condition, the need for terminological precisions makes it important
(for the further discussion) to observe that the complexity of a system, in mathematical
terms, refers not only to the number of components in the system but to the combination
of component-number and the strength with which the components are linked together
(connectance) (Ashby & Gardner 1970).

With respect to the type of equilibrium states of a system it is common, in mathematics

and its technology applications, to categorize them into equilibrium points and equilibrium
trajectories. In addition, the dynamic behaviour of the system as it strives (if at all) to return
to equilibrium, i.e., the extent to which, and the speed with which return occurs, can be
broadly studied in terms of resistance, elasticity, trajectory stability, and the extension of
potential stability domains. The use of the stability domain concept was first explored in relation
to diversity by May (1973, 1975), drawing on theories and techniques know as the Lyapunov
approach. We note that a system is seen as possessing higher stability in terms of resistance
the larger the force needed to displace the system from its equilibrium. With respect to
stability in terms of elasticity – in technology commonly also referred to as resilience – we
note that the faster the system returns to its equilibrium after a displacement, irrespective of

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 Bodin P. & Wiman B. L. B. – Resilience and other stability concepts in ecology

whether no, few, or many oscillations are involved, the more elastic (resilient) it is. The
importance of resilience in several cases outweighs that of other stability aspects; for instance,
the temporal response of a shock absorber is more important a property to consider than
the actual position of its stability (equilibrium) point. These concepts are illustrated in Fig.
1, depicting the concomitant behaviour of two components (“A” and “B”, for instance,
pressure and temperature in a fluid) resulting after a displacement of the system (for instance,
a boiler) from its equilibrium (or resulting from removing from the system a force that was
applied to maintain it at equilibrium).

Figure 1. Examples of differing types of equilibrium 2.

In (I), the system does have an equilibrium point but is basically unstable at this point – any
infinitesimally small displacement from that point results in a run-away behaviour of the
system. In (II), the system moves towards a stable equilibrium point, called focus. (III)
depicts a response to stress where the system moves without oscillations towards a node.
(IV) and (V) show that the equilibrium can be of a dynamic type (called centre) wherein
equilibrium is established by one or several trajectories. Case (VI), which is of basic concern,
shows that systems can possess what is known as stability domains (or “domains of attraction”)
within which the system, if displaced, can behave as in (II), (III), (IV), or (V), unless the
displacement force is large enough to bring the system to or outside the domain limits, in
which case a behaviour involving (I) occurs, and “surprise” might result.

Conceptual transfers to the ecology domain

Concepts such as those illustrated above can be tentatively moved into the domains of
ecology. The system can then be thought of as, for example, the exchange of energy or
biogeochemical substances between trophic levels (the components) in an ecosystem, or the
system for study can be conceived as an interplay between organisms. Assuming that the
system in itself does not change when subject to stress one can for instance attempt to
address the ecological system of one host (named A) and one parasite (named B) as a

2
Here, we have chosen to use the so-called phase-space for illustrations: co-variations among system
components along a time axis – the time-space – can be “condensed” through plotting the behaviour
of one component versus the concomitant behaviour of another component; time is then embedded
in the resulting phase-space plot such that the speed with which “movements” occur becomes visible
in, for instance, computer graphics. Figs. 2a and b illustrate the relationships between time-space and
phase-space.

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Bodin P. & Wiman B. L. B. – Resilience and other stability concepts in ecology

mathematical model where XA and XB represent the system variables (e.g. biomass, number
of individuals, energy content or carbon content) as follows:

dXA(t)/dt = fA(XA(t), XB(t)); XA(to)=XAo

Eq. 1
dXB(t)/dt = fB(XB(t), XB(t)); XB(to)=XBo

The solution to this set of differential equations can be plotted over time (Fig. 2a) or in the
phase-space (Fig. 2b) where the latter type of diagram in some respects shows the system
behaviour more clearly. Fig 2b shows how the system moves towards an equilibrium.

One common misconception is that an ecological system in equilibrium (such as “metabolic

equilibrium”; cf., e.g., Odum 1962, 1969, 1971) would be called “stable”. However, whether
or not a system is stable relates not to its being in equilibrium but to its capacity to remain
there; i.e., stability refers to the capability of the system to withstand forces tending to move
it away from equilibrium and to the behaviour of the system when, if displaced from
equilibrium, it strives to return (if at all) to equilibrium. Another not uncommon
misconception in ecology is that there exists a basic ability of systems to always return to
equilibrium, a kind of basic and (unfortunately) prevailing idea of ecological “homeostasis”
(also known as “the Laplacian demon”) that can be, and has been, challenged on several
grounds (cf. Wiman 1991).

Figure 2a. Hypothetical course of Figure 2b. The lapse with time in Figure a
development of a system of two co- plotted in phase space
varying species.

Early ideas about ecological dynamics – such as inherent in the Lotka-Volterra equations –
showed how populations theoretically could behave in a stable or an unstable manner (for
a review see e.g. Bodin 2004). Among the first to use the concept of resilience in the
ecological-systems realm was Holling (1973), who for that purpose partly abandoned the
traditional meaning of the concept in technology (see “elasticity” above), and re-defined it:
“Resilience determines the persistence of relationships within a system and is a measure of
the ability of these systems to absorb changes of state variables, driving variables, and
parameters, and still persist. In this definition resilience is the property of the system and
persistence or probability of extinction is the result.” (Holling 1973, p. 17) Thus, ecological
resilience as the ability of a system to return to equilibrium after a disturbance, and the

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 Bodin P. & Wiman B. L. B. – Resilience and other stability concepts in ecology

manner in which it would do so, and the several other implications of stability were
“condensed” into a new metaphor. In addition, community ecology, and theories of ecological
succession (cf., e.g., Odum 1969, Tilman 1999), have introduced partly differing perspectives
on stability (and its relationships with spatial scales, and with diversity; see below). This
means that several different meanings of stability co-exist, a state of affairs that to a significant
extent compounds the usefulness of analysis as well as synthesis (see e.g., Orians 1975,
Pimm 1984, Walker 1990).

Problems with chaotic behaviour in ecological systems

Holling’s (1978) ideas of stability and resilience were based on the observations of natural
systems showing multiple steady-states (cf. Ludwig et al. 1978). Catastrophic shifts –
incorporating various forms of hysteresis – in ecosystems have also been found and modelled
in several other ecological systems (Scheffer et al. 2001, see also Wiman 1991). This is one
example of (strongly) non-linear behaviour observed in ecological systems.

In the early 1970s Robert May (see Gleick 1987) studied the discrete form of the logistic
equation and observed that varying one variable could cause the simulated population to
regularly oscillate between two equilibria. Increasing the variable value further caused a
“period doubling” and yet larger values led to further period doublings and eventually to
completely chaotic behaviour (in terms of physical chaos, which is not to be confused with
“white noise”; cf. Stone 1998). In these debates on chaotic behaviour it soon turned out
that the relationships between stabilising/destabilising qualities and structural complexity
(such as diversity, connectance) can be of major importance (e.g., Ashby & Gardner 1970,
May 1972). For instance, Robert May investigated the supposed connection between stability
and diversity and found that in a Lotka-Volterra competition model, a greater diversity led
to lower local stability (Tilman 1999).

Stability and diversity in ecological systems

For a long time, diversity to ecologists seemed to be of little relevance to stability or other
population properties (Tilman 1999). However, as already indicated in the previous section,
the debate on stability in relation to diversity within ecology is long-standing (cf. Wiman
1991) and – partly because of prevailing misconceptions about diversity as well as stability –
the issue is far from resolved. In recent years, an increased interest in the old stability-
complexity debate has arisen. It has been suggested that increased diversity will lead to
increased stability, especially in the form of decreased variability (cf. Tilman 1996, 1999).
Some of these assumptions have been investigated and tested through a number of
simple models (Tilman 1999). A somewhat contradicting phenomenon that has been
observed is that an increased diversity would lead to decreased ecosystem variability but to
larger biomass variability at the individual population levels. According to Tilman (1996)
this observation can be supported by theory, and the existence of these types of scale-
dependent differences was in fact suggested in the early 1970s by May (1973). Some of the
confusion in the stability-complexity debate is probably due to a lack of communication
between community ecologists and population ecologists, because stabilising/destabilising
mechanisms very likely work differently at these different scales. In order to further develop

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Bodin P. & Wiman B. L. B. – Resilience and other stability concepts in ecology

theories and models of the diversity-stability concept, a bridging between the two sub-
disciplines of community and population ecology is needed (cf. Loreau 2000).

One problem with the whole stability-diversity debate (a full account of which is far beyond
the space available here) might reside in the several differing conceptions of equilibrium that
underpin analyses by differing scholars. As put by Loreau et al. (2001, p. 806): “[n]ew
approaches should be developed that take into account the dynamics of diversity and the
potential for adaptation through phenotypic plasticity, evolutionary changes and species
replacement”. As pointed out in the section about stability in the mathematics/technology
domain, particular challenges arise when trying to link fairly short-term ecological dynamics
to long-term evolution. With respect to the issue of system robustness and fragility, May
(1999) notes that ecosystems are in tension between evolutionary forces (that tend to add
species to efficiently exploit or subdivide every available niche) and dynamical forces (where
an increased species number leads to greater dynamical fragility).

It needs to be emphasised that the basic, stringent concepts of equilibrium essentially

emanate from models built on systems of ordinary differential equations that do not take
spatial structure into account. Thus, the equilibria reflect averages of population dynamics
parameters, and of ambient biotic and abiotic conditions (an approach known as “the
mean-field assumption in ecology”; cf. Dieckmann et al. 2000). Stochasticity added to these
models (in the population parameters or as added external stochastic forcings) tends to
make the populations become extinct (DeAngelis & Waterhouse 1987). Jansen and de Roos
(2000) address several aspects of how the dynamics and stochasticity of local (animal)
populations can be linked to give different large-scale (“global”) dynamics, including damped
oscillations on the large-scale spatial level (“statistical stabilisation”, i.e., oscillations in spatially
averaged densities become reduced because of reduction in oscillation amplitude in the
transition regions between local patches containing the populations, and because of averaging
in combination with phase differences in the oscillations occurring in the patches). To such
developments in spatio-temporal modelling, one should add the so-far seemingly little
explored potential that “[t]he distribution of ages within a population may be one of the
elements of diversity that contributes to the stability of the community – at least in the
sense that it permits or denies the chance of rapid recovery after a disaster” (Harper 1977, p.
707). This is, if “mean-field assumptions” are abandoned or at least complemented with
richness in space and age distributions, stability and diversity might well begin to take on
significantly more relevant meanings in ecology. As May (1999, p. 1954) puts the stability-
and-complexity issue: “In short, today’s conventional wisdom about the stability-complexity
relationship is vastly more textured than the ‘complexity begets stability’ of the 1960s.”

Bridges to sustainability science

Despite the fact that the transfer of the stability concepts (such as resilience) from the
mathematics/technology realm to ecology still is far from unproblematic, acceptance of
these concepts within the field has led to further transfers to other realms such as those
involving socio-economic dynamics. An example of such transfer is the “panarchy” concept
coined by Holling (2001) where an ecosystem holds one or more stable equilibria and, in the
process of succession, grows towards an increasingly productive but less and less stable (i.e.,
increasingly vulnerable) stage; finally, the increased strength of connection between the

38 the ESS Bulletin - Volume 2 Number 2 2004

 Bodin P. & Wiman B. L. B. – Resilience and other stability concepts in ecology

variables in the system leads to an abrupt change thus hypothesising that increased
complexity (sensu Ashby & Gardner 1970), yields decreased stability (Holling 1986). After
this change (caused by for instance fire, storms, pests, senescence), the ecosystem reorganizes
itself and the process of ecosystem renewal again commences. This process of growth and
abrupt shift is called an “adaptive cycle”. Such cycles are hypothesised (Holling 2001) to occur
at different time-scales (creating a hierarchy of adaptive cycles at different time-scales), and a
disturbance from a cycle on a shorter time-scale can cause a collapse at a longer time-scale.
Holling (2001) not only sees panarchy as a concept applicable to ecological systems but also
to economic and social systems, thus proposing that the concepts of resilience and stability
be taken a step further away from its original realms.

However, not only has the transfer of stability concepts from the technology to the ecology
realm proved problematic. In addition, along with the more precise definitions of diversity
that originate in information theory (based on statistical concepts of entropy; cf. e.g. Shannon
& Weaver 1949, Bell 1968, Gallager 1968) and in various ways are applied in ecology to reflect
species richness, species equitability and evenness, and complexity in the trophic differentiation
of food webs (cf. Walker 1990), several other diversity concepts are in use such as landscape
diversity, regional diversity, biogeographic diversity, and others (cf. Whittaker 1977). We
therefore prefer to adopt the stance that very limited progress has been made since May
professed that “[T]here is no comfortable theorem assuring that increased diversity and
complexity beget enhanced community stability; rather, as a mathematical generality, the
opposite is true. The task, then, is to elucidate the devious strategies which make for stability
in enduring natural systems. There will be no one simple answer to these questions” (May
1973, p. 174). As a consequence, we also recommend caution in the quest for further bridging
of resilience and diversity concepts from ecological to societal science realms.

Conclusions
As we have discussed above, the transfer of the stability concepts from the technology
realms to the ecological realms has not been a straightforward process and their definitions
within this new field are yet to be clarified. Still these concepts have further been transferred
into new fields moving further from their original definitions.

We argue that one ought to get back to the original definitions of stability and diversity
(complexity) in mathematics, wherein stability reflects the dynamics of a system in its
response to stress, and complexity is a measure of the number of components (in terms of
state variables) in the system and their connectance (strength of the linkages between these
components) (cf. Ashby & Gardner 1970). To these “basics” (in themselves far from trivial)
need to be added ongoing developments in spatio-temporal modelling and theory build-
up. Stabilising properties of the overall system might look very different if components
(populations) are modelled based on averages across space, in comparison to approaches
wherein responses in spatial sub-regions (“cells”) are first modelled and then linked together
to yield an aggregated community response.

We also observe that that at longer time-scales, when time-scales tend to evolutionary
measures, current ecological theory has very little to contribute. This is because over long
time-horizons, systems not only change, they also change how they change – and accounting

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Bodin P. & Wiman B. L. B. – Resilience and other stability concepts in ecology

for such phenomena in modelling faces severe challenges with respect to properly
incorporating changes in system structure itself in response to stress.

From a natural-resources management and precautionary-principle point of view, we argue

that it will offer best insurance against unmanageable surprise if the non-linearities involved
in ecological systems’ responses to stress be brought to the forefront. This would emphasise
the potential for stabilising properties being weakened in highly complex systems, in particular
if they are subjected to highly complex combinations of stress. That is, we essentially argue
the stance propounded by May in the early 1970s (cf. e.g. May 1975) that highly complex
systems should be perceived as having a larger potential for being fragile than do less
complex systems (although, assuming a sufficiently long time perspective, and a sufficiently
slow rate of change in stress factors, one might hypothesise that highly complex and fragile
systems might on the other hand hold a larger pool for structural change than less complex
and more robust systems).

Placing such a stance in a climate-change and society context can assist in advancing our
understanding of the extent to which ecosystemic shifts “spill over” to generate adverse
effects on societies that strongly depend on well-functioning landscapes and ecological
systems. An improved understanding of such chains of “spill-over effects” (from climatic
shifts to ecosystem shifts to effects on cultures and societies) can then provide support to
policy-making and “insurance-based” (i.e., risk-reducing) management of today’s and
tomorrow’s natural resources and their supporting ecological structures and functions.
While we do not propose to develop any kind of “predictive” methods to this end (because
of the potentially inherent non-linearities, even physically chaotic behaviour, in the above-
outlined chain of “spill-over effects”), we suggest that usable scenarios can result if the kind
of data exemplified above be combined with scenarios for climatic changes, for regions and
cases of particular importance.

We observe that already in the 1970s means were suggested – but so far little explored – by
which it is possible to introduce the judgment of the observer into stability analyses (cf.,
e.g., Harte 1979). This can be accomplished through invoking various mathematical
procedures which, for simplicity, can be named weighting. Such procedures can be used to let
the observer (such as a policy-maker) decide on which type of system-behaviour should be
judged as more important than another, and have the advantage of making a value-laden
assessment of a system behaviour transparent. Such an approach to combining technical
detail and stringency, in approaching ecosystem vulnerability and stability, with policy-relevant
interpretations and applications would at least place rigorous demands on clarity and precision
in defining the chain of concepts that underpin final decision-making.

Constraints and opportunities taken together, we recommend that any transfer of

technology-domain concepts into realms outside of technology – such as ecological,
behavioural or social sciences – would need as much careful scrutiny as would the transfer of
concepts from for instance social science into the domain of technology or ecology. Any
such deeper “transfer analysis” is beyond the scope of this contribution, and we expect
analyses of that kind to become one salient topic for discourse during the Symposium
workshop deliberations.

40 the ESS Bulletin - Volume 2 Number 2 2004

 Bodin P. & Wiman B. L. B. – Resilience and other stability concepts in ecology

Acknowledgements
We thank Göran I. Ågren for his constructive criticism of an earlier version of this
contribution.

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