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journal homepage: www.elsevier.com/locate/ecolmodel

Resilience in landscape exploitation systems

C.S. Fletcher , David W. Hilbert
Tropical Forest Research Centre, CSIRO Sustainable Ecosystems, PO Box 780, Atherton, Queensland 4883, Australia

a r t i c l e

i n f o

a b s t r a c t

Article history:

A generic model is developed that describes the broad properties of land-exploitation sys-

Received 16 March 2006

tems from hunting-gathering societies to modern, intensive agriculture. The framework

Received in revised form

includes social, economic and ecological drivers of change, going beyond the concept of

13 October 2006

sustainability to establish a paradigm of resilient exploitation that calculates the capac-

Accepted 17 October 2006

ity of landscape exploitation systems to survive in uncertain and variable human and natural
environments. The model is highly aggregated, consisting of two state variables: (1) humanmade capital and labour (H) and (2) natural capital (N). Depending on the parameters, the

Keywords:

model displays a single non-trivial equilibrium, two equilibria, or stable limit cycles. Our

Ecological economics

analysis focuses on the strategies that exploiters employ, through varying their investment

Natural capital

in H, and how this affects resilience of the system. We measure resilience as the size of

Predatorprey model

the basin of attraction near a desirable equilibrium and the return time following small

Grazing systems

perturbations. Four general strategies are analysed and discussed using grazing systems as

Sustainability

a particular example: (1) constant stocking rate, (2) maintaining grass stock, (3) constant

Agriculture

utilization rate, and (4) a non-linear, compound strategy.

Like previous models of renewable resources, maximum sustainable yield and prot are
determined by properties of the renewable resource, so all management strategies yield
identical long-term sustainable production and identical equilibria. However, different management strategies drastically change the systems resilience. We assess the opportunity
cost of the disparate objectives of resilience and protability using multi-objective optimization and demonstrate that resilience decreases rapidly as maximum prot is approached.
Thus, the same stable and sustainable equilibrium point in an exploitation system will be
more or less resilient depending on the exploiters strategy. Traditional economic analysis
does not consider the impact of the management strategy employed to drive the system
towards these long-term equilibria, and therefore misses large differences in dynamical
behaviour away from equilibrium. The possibility of multiple steady states in these nonlinear systems results in the potential for a discontinuous threshold in resilience, to one
side of which resilience is maintained, while on the other resilience and production are
greatly degraded. We further discuss how subsidies might increase or decrease resilience of
exploitation systems by affecting an exploiters strategy.
2006 Elsevier B.V. All rights reserved.

. . . proactive action to manage systems sustainably and

to build resilience into systems will be advantageous, particularly when conditions are changing rapidly, when surprise events are likely, or when uncertainty is high . . .
Millennium Ecosystem Assessment (2005).

1.

Introduction

World-wide, human beings exploit natural capital for their

economic and social well-being. By doing so they create com-

Corresponding author. Tel.: +61 7 40918820; fax: +61 7 40918888.

E-mail address: Cameron.Fletcher@csiro.au (C.S. Fletcher).
0304-3800/$ see front matter 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.ecolmodel.2006.10.011
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plex systems with interacting economic, social and ecological

sub-systems. However, experience suggests that humancreated land-exploitation systems often fail in the long run
(Brander and Taylor, 1998; Folke et al., 2004; Millennium
Ecosystem Assessment, 2005). This raises important and
practical questions. Are these unsuccessful systems simply
overexploited to collapse, or do they suffer a more subtle
gradual erosion of resilience to natural environmental variability until collapse is random but inevitable? Is it possible
to identify short and medium-term management behaviours
that can increase the long-term resilience of a system while
still remaining protable?
There are examples of institutional arrangements designed
to achieve multiple long-term and socially optimal goals
(Ostrom, 1990). More commonly, however, humans have
demonstrated that they can most efciently optimize the
performance of their systems for a single, easily identiable goal, such as prot, as long as there is a strong, fast
feedback between management strategy and system performance (Clarke, 1990). Historically, this may have selected
management strategies with low overhead costs and efcient operation, with no impetus to select strategies that
foster real insight into the long-term operation of these
systems.
This is not a good management framework in the longterm because short-term tactics break down when (1) systems exhibit highly non-linear response, such as threshold
effects (Muradian, 2001); (2) when an exploiter wants to optimize across multiple objectives which may not combine intuitively or linearly (Romero and Rehman, 1989; Zander and

Kachele,
1999); or (3) when the behaviour to be optimized is
not an easily and rapidly measured physical quantity (Ives,
1995). Each of these situations is becoming more important as intensication of humanecological systems increases
(Millennium Ecosystem Assessment, 2005). We now know
that non-linear thresholds often exist in landscape exploitation systems (Muradian, 2001; Scheffer and Carpenter, 2003)
and that long-term persistence requires management strategies that balance short and long-term economic gain with
less tangible indicators, such as resilience (Folke et al., 2004).
Resilience is a vital extension to the concept of sustainability (Walker et al., 2004), because it embraces the importance
of environmental variability and stochastic shocks, which
determine the long-term persistence of complex systems,
especially climate-forced, natural systems (Mangel and Tier,
1994).
Serendipitously, humanitys capacity for understanding
and exible management provides the opportunity to optimize systems for something more than just maximum productive or reproductive output (Hardin, 1968), as long as
information exists to support management decisions. Being
aware of the opportunity cost implied by different management strategies in terms of system resilience would allow
managers to make a conscious trade-off between prot
and resilience, possibly ameliorating many of the negative consequences currently observed when humans modify natural ecosystems (Millennium Ecosystem Assessment,
2005).
We begin to address these issues for landscape exploitation systems with a multi-objective optimization analysis of a

dynamical systems model that captures the social, economic

and ecological interactions that are important in these coupled systems. Landscape exploitation systems arise when an
exploiter, or a group of exploiters operating cooperatively to
apply a consistent management strategy, chooses to employ
a stock of human-made capital to extract part of a renewable stock of natural capital from the local environment to
create a desirable economic good, the consumption or sale
of which nances increased reinvestment in new humanmade capital. This closed system, in which the growth of
human-made capital is related to its consumption of natural capital, is reminiscent of the classic predatorprey model
of ecology introduced by Lotka and Volterra (Volterra, 1931;
Lotka, 1932). The distinction, however, is that the system is
managed by an exploiter who applies management strategies to achieve their personal goals. It is the existence of the
manager that ties the environmental and ecological parts of
an exploitation system to human economic and social systems. This creates complicated interactions whereby exploitation of renewable natural resources drives the economic
and social development of society, and the demands and
expectations of society effect how the exploitation system is
managed.
This denition is intentionally broad enough to cover
a variety of systems; from hunting-gathering and swidden
agriculture systems, to grazing and modern intensive agriculture systems. This paper investigates all generic landscape exploitation systems simultaneously, by characterizing, for example, a grazing system in the same manner as
a hunting-gathering system, by assuming human-made capital
(e.g., domesticated plants and animals or humans) consumes
natural capital (e.g., grass or bush meat) at a certain rate, to
produce an economic good (e.g., beef or nutritional food). We
characterize exploitation strategies in the same abstract manner: a grazier or a hunter and gatherer might believe that
the future production capacity and protability of their system is primarily a function of their total stock of humanmade capital, or the quality and quantity of their natural capital, or perhaps a balance of the two. The understanding of
how their system works will effect the management decisions that an exploiter makes, and change the long-term
dynamics of the exploitation system. Interestingly, it will
not change the maximum productive capacity of the system, because that is fundamentally limited by the growth
rate of natural capital and the way in which a given type
of human-made capital processes natural capital (Clarke,
1990). These processes can be quantied using the tools of
dynamical systems theory to calculate both long-term prot
and resilience, which can be analysed using multi-objective
optimization to visualize the trade-off that an exploiter
must make when trying to balance these two management
objectives.
Simple mathematical models of specic exploitation systems have a rich history, including models of Easter Island
(Brander and Taylor, 1998), the Anasazi (Janssen et al., 2003),
grazing systems models (Noy-Meir, 1975; Anderies et al., 2002),
and the early sheries models that lead to the maximum sustainable yield (MSY) paradigm (Gordon, 1954; Schaefer, 1957).
Models of generic economicsocialecological systems have
also been published, such as studies of technology-fuelled

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Table 1 Model parameters, their units, denitions, and simulation values

Parameter

Value

r
K
Cmax


1
100
2
0.5

b


0.05
N/A

Description and units

Maximum growth rate of natural capital (time1 )
Carrying capacity of N, i.e. amount of N in a stable unexploited system (N)
Maximum consumption rate of natural capital per unit of human-made capital (N H1 time1 )
Half-saturation of decreasing marginal returns, i.e. time required to consume half remaining natural capital
at maximum consumption rate (time)
Production coefcient, i.e. how consumed N is transformed into production, measured relative to the value
of a unit of H (H N1 )
Rate at which human-made capital depreciates (time1 )
Savings rate: fraction of production that is reinvested to purchase new human-made capital (unitless)

economic and population growth relying on a limited natural

resource base (Anderies, 2003), and more complicated simulation models (Woodwell, 1998). These models have often been
applied to the rise and fall of civilization, economic or population growth, or the capacity for technological change to
overcome resource limitations.
These studies often focus on either specic dynamics, such
as collapse trajectories of early civilizations, or generalized
static analyses, such as estimates of maximum sustainable
yield. In contrast, our model framework aims to capture general dynamic information, using summary indicators such as
resilience, to calculate the capacity of a system to avoid any
trajectory leading to collapse in an uncertain and variable
environment. We ask the question: if an exploiter approaches
their exploitation system with a given type of mind set, and
goals driven by given social values, what is the likely outcome,
in an environment of given natural richness, over medium
and long-term time-scales? The results apply from huntinggathering societies through to modern intensive agriculture,
allowing us to identify fundamental social, economic and ecological drivers of resilience across all landscape exploitation
systems.

2.

The model

The model is similar to the canonical predatorprey system in

which human-made capital, H, consumes natural capital, N.
These variables are aggregate measures of sub-systems which
may be quite complex in detail. Land area is assumed xed
and sufciently homogeneous that spatial variability can be
ignored.
Natural capital grows logistically at an intrinsic rate r with
a carrying capacity K. It is consumed by human-made capital at a maximum rate of Cmax (per unit of H), modied by an
efciency factor between 0 and 1 determined by the functional response, f[N,H], which is dependent on the stocks of
both types of capital. These relations are shown in the following equation:

= rN 1
N

N
K

f [N, H]Cmax H

(1)

Parameter denitions, units and values used to generate gures are summarized in Table 1. The amount of economic good produced (P) is directly proportional to the
amount of natural capital consumed, with a proportionality

constant a:
P = af [N, H]Cmax H

(2)

Human-made capital may be increased by reinvesting production into the system, using a fractional savings rate, [N,H],
rather than extracting prot to invest in alternatives available in the wider economy. In the absence of reinvestment,
H depreciates exponentially at a rate, b. Thus, the dynamical
equation for H is
= [N, H]P bH = [N, H]aCmax f [N, H]H bH
H

(3)

The functional response:

f [N, H] =

N
Cmax H + N

(4)

describes the efciency with which a unit of human-made

capital consumes natural capital, relative to its maximum possible consumption rate, Cmax .
This particular form can be interpreted as both a decreasing
marginal return to increased investment in human-made capital at a given level of natural capital, illustrated in Fig. 1(a), and
a weak-complementarity between the natural and humanmade factors of production (Kraev, 2002), as illustrated by the
isoquant diagram in Fig. 1(b). Fig. 1(a) shows how  parameterizes this relationship: the point at which decreasing marginal
returns reaches half-efciency is H = N/Cmax .
All exploitation systems exhibit some fundamental level
of decreasing marginal returns due to increasing difculty of
harvesting marginal resources (Tainter, 1998), but it is also possible for an environmentally conservative exploiter to choose
an articially high value of  in order to safeguard their system
against a transient surplus of H. In this context, the functional
response represents a self-imposed temporary reduction in
consumption and production efciency that an exploiter uses
to prevent over-exploitation of their natural capital base. As
a variable, shrinking to maximize prot at the expense of
reduced resilience during times of constancy, and growing to
maximize resilience at the cost of prot during times of variability, it might be a proxy for adaptability (Walker et al., 2004).
In contrast to this short-term and inefcient emergency
management for unbalanced systems, it is manipulation of
the savings rate that represents the primary way an exploiter
manages their system over the long-term to balance the levels
of natural and human-made capital to maintain efciency in
an effort to achieve their performance goals. To do this they

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Fig. 1 (a) shows that in a system abundant in natural capital, N  Cmax H, consumption efciency will be nearly
maximized, f[N,H] 1, and relatively small increases in human-made capital will exhibit almost-constant marginal returns.
However, in a system where the amount of natural capital consumed by human-made capital is already comparable to or
greater than the amount of natural capital available, N Cmax H, increased investment in H will yield rapidly decreasing
marginal returns. This complementarity reects the fact that human-made and natural capital are only weakly
substitutable; both are vital to the consumption and production processes of the exploitation system, and increasing H
when the system is limited by N is not an efcient strategy (Costanza and Daly, 1992; Kraev, 2002), as indicated by the fact
that the isoquants in (b) asymptote to a nite value away from the axes. On the other hand, an exploiter will be nancially
rewarded for reinvestment as long as the marginal return indicated in (a) is greater than that of alternative investments.
Parameter values are as shown in Table 1, with the exception that Cmax = 5.

have to make decisions about issues like whether their system

is operating sustainably and whether it will reward reinvestment at a higher rate than alternative investments.
We imagine that an exploiter makes these decisions based
on a simplied mental model of their exploitation system, which describes how an exploiter expects their system to respond to management decisions. Fundamentally,
the exploiter is trying to estimate the future productivity of
their system under a given reinvestment strategy to calculate whether the return on such a reinvestment would exceed
current investment market alternatives. The mental model by
which an exploiter understands the capacity of their system
to produce a prot is a function of many factors, including
personal drivers such as their own experience of their system,
social drivers such as the exploiters education and the norms
of the society in which they operate, and economic drivers,
such as taxes and subsidies.

3.

Analysis

To generate and analyse a specic model, we must specify

a mental model or savings rate function representing the
management strategy of the exploiter. We investigate four
simple but qualitatively distinct exploitation strategies that
reproduce many dynamic behaviours appropriate to landscape exploitation systems, such as sudden collapse, cyclic
behaviour, and long-term stability. We avoid analysing this
plethora of specic trajectories by analysing the generic
dynamic properties of these systems, such as their resilience,
and more traditional measures of performance, such as profitability. We optimize the system across both of these objectives simultaneously to illustrate the compromise faced by
an exploiter trying to attain both reasonable economic and
acceptable dynamic performance. This analysis goes beyond
traditional steady state optimization by intentionally studying
non-economically optimal solutions to provide information

about the opportunity cost of economically rational management in a variable natural environment.
Although we choose both specic systems and specic
quantitative analyses in order to perform numeric calculations, for the purposes of this paper we are more interested
in characterizing the types of model behaviour, and types of
analysis possible using our framework. These examples represent a starting point that can be adapted to more realistic
situations.
Here, we interpret each strategy within the context of a
grazing system, before generalizing the results to all exploitation systems. We examine four qualitatively distinct exploitation strategies: managing for constant stocking rate; managing
to maintain a constant grass stock; managing to achieve constant utilization by balancing the ratio of cattle to grass; and
a compound strategy combining aspects of the other three at
different levels of abundance. These strategies modify how
an exploiter reinvests to build human-made capital in their
system, and consequently each strategy exhibits a distinct H
growth equation. In contrast, each strategy exhibits the same
N growth equation, reecting the fundamental limits placed
on exploitation systems by their natural capital base.
This information is represented graphically in the isocline
plots of Figs. 25(a). As the natural capital growth rate is independent of exploitation strategy, the same N isocline:
H=

rN(N K)
Cmax (r(N K) + K)

(5)

is used throughout each analysis. In comparison to the classic inverted parabola of traditional predatorprey models, this
isocline exhibits a skewed peak Npeak < K/2 due to diminishing
marginal returns. The H isocline varies between management
strategies, but the mathematical forms of each strategy have
been chosen to provide a range of simple and qualitatively
distinct isoclines that conveniently summarize possible system behaviours. For the sake of simplicity, the H isoclines

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Fig. 2 Maintaining a stock of human-made capital. (a) shows the N (faint line) and H (heavy line) system isoclines, with
Hgoal such that the system maximizes prot, superimposed on a faint vector eld showing the direction of change at each
point in state space. The desirable stable point occurs at the right-most intersection of the isoclines, the turning point at the
left-most intersection of the isoclines, and the undesirable stable point at the intersection of the H isocline and the vertical
axis. (b) shows the real (solid) and imaginary (dashed) values of the two eigenvalues of the linearized system (light grey,
dark grey), indicating that the desirable critical point is asymptotically stable across the whole management region. (c) plots
economic indicators vs. the control parameter accessible to the exploiter; (d) plots dynamic performance across the same
region; and (e) plots the Pareto front generated by a multi-objective optimization analysis of the system across the dual
objectives of prot and resilience.

are not constrained to yield a savings rate function [N,H]

bounded below [N,H] = 1, implying that an exploiter can reinvest to build their system faster than they generate prots,
perhaps using credit. It is possible to constrain the savings
rate function appropriately with a more complicated function, and the steady state results presented herein remain the
same.
Figs. 25(a) illustrate an N isocline and an H isocline that
contain the points in state space where each state variable
exhibits zero growth. In the region to one side of an isocline
the dynamics driven by natural processes and management
actions act to reduce the relevant state variable; while in the
other region the state variable increases. Only along the iso-

cline is a state variable in equilibrium. At the intersection of

the N and H isoclines, both state variables are in equilibrium
and the system occupies a steady state, which is important
because it determines the long-term performance of the system. However, the critical points, and key measures of statespace topology about the critical points, also represent our
best summary of the general dynamics of systems, including
stochastically driven systems that may never actually attain
the long-term steady state.
The nature of the desirable critical point is illustrated in
Figs. 25(b), using the eigenvalues of the system linearized
about the critical point. When both eigenvalues are purely
real and negative, the desirable critical point is asymptotically

Fig. 3 Maintaining a stock of natural capital. (a) shows the N (faint line) and H (heavy line) system isoclines, with Ngoal
such that the system maximizes prot, superimposed on a faint vector eld showing the direction of change at each point
in state space. The desirable critical point occurs at the intersection of the isoclines. (b) shows the real (solid) and imaginary
(dashed) values of the two eigenvalues of the linearized system (light grey, dark grey), indicating that the desirable critical
point is asymptotically stable for Ncrit [95,100], oscillatory and stable for Ncrit [50,95], unstable supporting a stable limit
cycle for Ncrit [30,50] and unstable for Ncrit [0,30]. (c) plots economic indicators vs. the control parameter across the range
of stable solutions Ncrit [50,100]; (d) plots dynamic performance across the same region; and (e) plots the Pareto front
generated by a multi-objective optimization analysis of the system across the dual objectives of prot and resilience.
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Fig. 4 Balancing stocks of human-made capital to available natural capital. (a) shows the N (faint line) and H (heavy line)
system isoclines, with Rgoal such that the system maximizes prot, superimposed on a faint vector eld showing the
direction of change at each point in state space. The desirable critical point occurs at the intersection of the isoclines. (b)
shows the real (solid) and imaginary (dashed) values of the two eigenvalues of the linearized system (light grey, dark grey),
indicating that the desirable critical point is asymptotically stable for Ncrit [95,100], oscillatory and stable for Ncrit [40,95],
unstable supporting a stable limit cycle for Ncrit [10,40] and unstable for Ncrit [0,10]. (c) plots economic indicators vs. the
control parameter across the range of stable solutions Ncrit [50,100]; (d) plots dynamic performance across the same region;
and (e) plots the Pareto front generated by a multi-objective optimization analysis of the system across the dual objectives of
prot and resilience.

stable. When the eigenvalues are complex conjugates with

negative real parts the critical point exhibits oscillatory stability. If the system is modied, moving the critical point such
that the real part of the conjugate eigenvalues passes through
zero to become positive, the system undergoes a Hopf bifurcation and the critical point becomes unstable, but supports a
stable limit cycle. When both eigenvalues are purely real and
positive, the critical point is an unstable node. The economic
and dynamic properties of the desirable critical point are
calculated across the region Npeak < Ncrit < K, where the critical
point is stable for every management strategy.

Two measures of economic performance are calculated:

total prot, and marginal prot to investment in human capital. Prot is strongly related to total production, but also
depends explicitly on the savings rate strategy:

 = (1 [N, H])P = (1 [N, H])a

Cmax NH
Cmax H + N

(6)

Marginal prot is the derivative of prot with respect to

human-made capital. It is a useful economic measure because
it illustrates the point at which economic compulsion to inten-

Fig. 5 A compound strategy. (a) shows the N (faint line) and H (heavy line) system isoclines, with Sgoal such that the
system maximizes prot, superimposed on a faint vector eld showing the direction of change at each point in state space.
The desirable stable point occurs at the right-most intersection of the isoclines, the turning point at the middle intersection
of the isoclines, and the undesirable stable point at the left-most intersection of the isoclines, but for different values of Sgoal
it is possible that only the right-most or left-most critical point exists. (b) shows the real (solid) and imaginary (dashed)
values of the two eigenvalues of the linearized system (light grey, dark grey), indicating that the desirable critical point is
asymptotically stable across the whole plot region. (c) plots economic indicators vs. the control parameter accessible to the
exploiter, showing the protability of multiple steady states when they exist; (d) plots dynamic performance across the
same region; and (e) plots the Pareto front generated by a multi-objective optimization analysis of the system across the
dual objectives of prot (of the desirable steady state) and resilience.
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sify a system is lost. However, it explicitly values investment in

human-made capital while not assigning a value to stocks of
natural capital, which is arguably a major drawback of modern
economic systems, and a primary area of study for our model
(Costanza and Daly, 1987, 1992; Costanza et al., 1997).
We consider two indicators of dynamic performance
(Holling, 1973): (1) the resilience, or size of the basin of attraction, refers to the distance along the N axis between the desirable attractor (uppermost and rightmost attractor) and either
N = 0, in the case of a system with only one non-trivial attractor, or the non-trivial unstable node, in the case where there
is more than one non-trivial critical point in the system; and
(2) return time required to recover following a small perturbation away from a stable point (May et al., 1974; Beddington
et al., 1976). This is a useful indicator of dynamic performance
because a stochastically forced system with a long return time
will exhibit large statistical variance about the deterministic mean, effectively reducing the resilience of the system
to repeated perturbations (Beddington and May, 1977). Return
time in our analysis is based on the standard denition: the
negative and inverse value of the larger of the real parts of the
two eigenvalues of the linearized system about the desirable
critical point in the system (Beddington et al., 1976).
For each management strategy, estimates of the economic
and dynamical properties of the steady state of the system
are calculated across the range Npeak < Ncrit < K, where every
strategy exhibits at least one stable critical node. That is, we
do not assume that an exploiter automatically aims to optimize the economic performance of their system as part of our
model structure. In this way, we can optimize for a combination of objectives including both economic performance and
resilience; and assess the opportunity cost of any particular
combination of these parameters.
Multi-objective optimization provides information about
the trade-off between optimizing various objectives. We optimize prot and resilience to illustrate the fundamental tradeoffs and opportunity costs for each management strategy. We
use an adaptive MOGRASP algorithm developed by Higgins
et al. (2006). The Pareto front is generated from the nondominated solutions to the set of optima derived from different, adaptively chosen weightings of the combined objective
function of the form:
Objective = w Prot + (1 w) Resilience,

w [0, 1]

(7)

Practically, the Pareto front represents the best possible

resilience accessible at each value of prot, and vice-versa.

3.1.

Constant stocking rate

Traditional economic theory emphasizes the importance of

human-made capital as the limiting factor of production in
exploitation systems, under the assumption of abundant natural resources (Foster, 2003). If a grazier, for example, believed
that prot was primarily determined by the total head of cattle they ran, they might aim to maintain some desired stock
of cattle, Hgoal , independent of the state of the natural capital of their system. Traditional economic theory may expect
an exploiter to set Hgoal to a value that matched expected
marginal returns to that available from alternative invest-

ments, but the exploiters personal preferences can also affect

their choice. Instead of specifying how the exploiter chooses
Hgoal , we calculate the economic and dynamic performance
for various values of Hgoal , and analyse the trade-offs.
Such a system will exhibit a horizontal H isocline like that
in Fig. 2(a), based on a human-made capital growth rate:
= (Hgoal H)H
H

(8)

This growth rate can be related to that in Eq. (3) by specifying an appropriate savings rate function, [N,H], which represents how the exploiter would choose to reinvest based on
their mental model. Fig. 2 and its caption summarize the systems quantitative behaviour in terms of critical points and
their stability.
The shaded vector eld shows how trajectories from many
initial conditions fall into two basins of attraction, evolving
towards the desirable critical point in Npeak < Ncrit < K and the
undesirable critical point at (0,Hgoal ), delineated by a separatrix curve passing through the unstable critical point in
0 < Ncrit < Npeak , where Npeak is the value of N for which the
peak in the N isocline occurs (constant across all management
strategies), and Ncrit is the (variable) value of the position of
the desirable critical point. From the exploiters perspective,
it is important not to let the system move from the desirable
basin of attraction across the separatrix, even as a result of
stochastic forces outside their control. Note that a disturbance
or sequence of small disturbances that moves the system
across the separatrix may leave a signicant amount of forage indicating a damaged but healthy system, encouraging
the exploiter to continue business-as-usual, a tactic that will
drive the system to eventual collapse at (0,Hgoal ). This is the
importance of resilience as a measure of how big a shock a
system can absorb before it undergoes invisible fundamental
structural change leading to degradation or collapse in a nonintuitive, non-linear way.
In this scenario, the exploiters choice of Hgoal is the primary management strategy. Fig. 2(c) shows how the desirable steady state values of the two economic indicators vary
with the choice of Hgoal . A value of Hgoal 17 maximizes prot
at approximately 24 H-equivalent units over the time scale
set by units of the maximum consumption rate. This corresponds to an H isocline that intersects the N isocline close to
its peak, but it also reduces resilience (basin size), as described
below. Marginal prot is maximized at minimum Hgoal , and
decreases monotonically as Hgoal increases, reecting diminishing marginal returns as Hgoal increases and Hcrit decreases.
Economic theory would predicate operating at the lowest
prot-per-unit-H that still exceeded marginal returns of alternative investments. That is, if the grazier could receive, say
10% interest on a secure term deposit, economic motivation
for exploitation would be lost at Hgoal = 16.6 when the marginal
benet dropped below this return.
The dynamic properties of the system are illustrated in
Fig. 2(d). The key result is that resilience decreases increasingly
rapidly from its maximum value in an unexploited system to
zero as intensication increases, Hgoal Hpeak . Return time is
minimized at Hgoal = 16.4, and it is comparable to the other
strategies across most of the management region.

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The conundrum faced by a grazier worried about both economic and dynamic performance is what choice of Hgoal to
apply, given that economic and dynamic performance are optimized at very different values. This is particularly apparent in
the conict that exists between the MSY paradigm and our
measures of resilience. MSY calculates the maximum yield
that the system can sustain indenitely, but explicitly ignores
the possibility of unexpected disturbances. All of the steady
state points forming the economic performance curves in
Fig. 2(b) are sustainable, and MSY occurs for maximum production, Hgoal 17. Resilience, in contrast, explicitly addresses
the capacity of the system to absorb shocks, and in our analysis yields the result that the minimally exploited system
maximizes this capacity, at Hgoal 0. Accepting that exploitation systems are an important part of human systems, so
that some compromise between production and permanence
must be reached, there is useful information in the form
of the trade-off between maximizing yield and maximizing
resilience.
Fig. 2(e) illustrates the trade-off between prot and
resilience in this system as a Pareto front (Miettinen, 1998).
Our model leads to a fairly simple Pareto structure in the
protresilience objective space. Interestingly, the slope of the
front is relatively consistent across the left 75% of the curve,
but steepens sharply toward the right. This implies that, for an
exploiter at position A on the front, a move to B in an effort
to extract the last 25% of economic performance degrades the
remaining resilience by approximately 70%.
The minimum resilience indicated on the Pareto front is
not zero, despite the fact that this system can exhibit zero
resilience, because there is a peak in the protability of the
system away from the absolute minimum resilience state,
and beyond this peak there is no economic compulsion to
decrease the resilience of the system. Fundamentally, this
arises from the skew of the N isocline: a system with constant
marginal returns would encourage exploitation to a point
close to zero resilience and poor stability (Beddington and May,
1977).
Although we have referred to a grazing system interpretation throughout, the model applies to all exploitation systems, from hunting and gathering societies to modern intensive agriculture. In this sense the form of the model represents an exploiter who sees natural capital as abundant and
believes the capacity of their system to yield protable returns
on further investment is determined by human-made capital alone. Such a system will exhibit two basins of attraction, implying a strong trade-off between maximum prot and
resilience.

3.2.

Maintaining grass stock

If a grazier believed that the protability and long-term viability of their property was primarily dependent on their stock
of natural capital, they might choose to apply a management
strategy aiming to maintain a constant stock of natural capital, Ngoal , by adjusting H while considering alternative investments that may promise more favourable rates of return.
Implicitly, the grazier would manage their ranch to operate at
N = Ngoal , in anticipation of the greatest long-term sustainable
performance of their system.

Such a system will exhibit a vertical H isocline like that

illustrated in Fig. 3(a), based on a human-made capital growth
rate:
= (N Ngoal )H
H

(9)

In contrast to the previous case, there is only a single basin

of attraction about the desirable critical point at the intersection of the isoclines. Across the parameter regions that we
study below, this simple state-space topology implies that any
grazing system with non-zero quantities of both cattle and
grass will eventually reach the desired critical point, as indicated by the faint vector ow eld in Fig. 3(a). Dynamically, this
means that perturbations would have to completely eliminate
the stock of either cattle or grass before the system exhibited
a qualitatively different dynamical behaviour over long timescales. This more robust long-term dynamic performance is a
result of the exploiters willingness to react to an unexpected
loss of natural capital by temporarily curtailing short-term
production to allow the system to recover. However, this reactive strategy is at risk of collapse in a stochastic environment
with repeated or coincident events.
Changing Ngoal varies the feedback between growth of natural capital and its consumption and reinvestment in new
human-made capital, changing the stability of the desirable
critical point as indicated in Fig. 3. We limit the following
analysis to the region, Npeak < Ngoal < K, where the system is
stable because exploiters are unlikely to desire a cyclical system unless several local systems are exploited out of phase,
such as a swidden agricultural system.
The exploiters choice of Ngoal leads to varying economic
and dynamic performance, shown in Fig. 3(c) and (d), respectively. A value of Ngoal 50 maximizes prot. Marginal prot is
maximized at maximum Ngoal , which corresponds to minimizing the amount of H in the steady state, reecting diminishing
marginal returns.
The return time across the region Npeak < Ngoal < K is minimized at a value of Ngoal 85. Resilience is maximized for
an unexploited system (Ngoal = 100) and decreases with any
increase in exploitation. It is minimized at the peak in the
N isocline, when the system can survive at most a reduction
of 41.4 units of N without structural change. This reects the
fact that at the very edge of our chosen management region
the system does not suddenly collapse, it merely takes on
less desirable dynamic characteristics (stable limit cycles),
in contrast to the previous example system. This is an important distinction, because in this system a perturbation out of
the desirable region will not lead to an almost-irreversible
change in system dynamics.
The trade-off between economic and dynamic performance is indicated as a Pareto front in Fig. 3(e). Similar to the
previous case, achieving the last 25% economic performance
implies an increasingly rapid decrease in resilience. However,
because the minimum resilience is not zero, the system maintains a signicant buffer even when it is being operated at
maximum prot.
This analysis applies equally well to all types of exploitation system. This is perhaps the simplest strategy consistent
with a manager that recognizes the limitation of natural capital as the signicant factor in production.

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3.3.

responsivity, Sgoal :

Constant utilization rate

Possibly a grazier recognizes the importance of balancing the

number of cattle to the current grass stock, so that they aim
to maintain a goal ratio of cows to grass, Rgoal = N/H. Such a
system will exhibit an H isocline linear in N like that in Fig. 4(a),
due to the H growth rate:
= (Rgoal N H)H
H

(10)

Similar to the last case, this system exhibits a single basin

of attraction centred about the critical point at the intersection
of the N and H isoclines, as long as the exploiters goal ratio,
Rgoal , is smaller than the gradient of the N isocline at the origin.
As in the rst scenario, the system exhibits stable limit cycles.
We limit our analysis across the management region to the
right of the peak in the N isocline, 41.4 N 100, where the
system always exhibits a stable node.
For a given exploitation system in a given environment, it is the exploiters choice of Rgoal that forms the
primary management strategy. Fig. 4(c) shows how the
steady state values of the economic indicators vary with
the choice of Rgoal . A value of Rgoal 0.33 maximizes prot.
Marginal prot is maximized at minimum Rgoal , which corresponds to minimizing the amount of H in the steady
state.
The dynamic properties of the system are illustrated in
Fig. 4(e). The return time is presented across the region
Npeak < Ngoal < K, and is minimized at a value of Rgoal 0.1. The
resilience is optimized at the edge of the control region at
Rgoal 0. Again, the minimum value of resilience at N = 41.4
is not zero, and the system can recover from a maximum perturbation of 41.4 units of N.
This strategy is similar to the previous case in that prot
and resilience both vary in Ncrit (not plotted) identically to
the previous case, although return time differs. Importantly,
this does not imply that the systems are identical from
the exploiters perspective, because each exploiter interacts
with their systems primarily through their choice of control
parameter. One obvious result is that an exploiter choosing the
current strategy faces a small absolute change in their control
parameter for large changes in system behaviour, while a grazier managing for forage stock faces small relative changes
in their control parameter. This has implications depending
on the accuracy of the exploiters model and their ability
to measure their chosen control parameters quickly and
regularly.

3.4.

A compound strategy

In general, an exploiter can adopt an arbitrarily complicated

management strategy, including one in which they change
their strategy in response to the severity of a perturbation.
One way of beginning to capture this process is by describing a compound strategy that switches between each of the
three previous strategies, depending on the balance between
cows and grass in the system. Such a strategy could exhibit
a saturating H isocline, shown in Eq. (11) and Fig. 5(a), with
an exploiter-chosen: maximum number of cattle, H = S2 ; minimum amount of grass for viable exploitation, N = S1 ; and

=
H

Sgoal

N S1
H H
N + S2

(11)

This produces a compound strategy as illustrated in

Fig. 5(a), or can qualitatively reproduce any of the previous
exploitation strategies, depending on the choice of strategy
parameters relative to the natural and economic parameters
of the model.
A compound strategy can generate a signicantly more
complicated state-space topology than the simpler cases previously considered. Depending on the exploiters choice of
control parameter, the system may exhibit a single basin of
attraction about either the left-most or right-most intersection of the isoclines, or two basins of attraction about both
these points, delineated by a separatrix passing through the
middle intersection of the isoclines. Fig. 5(a) shows the system
in a two-basin conguration, as illustrated by the vector ow
eld.
The most important observation about this strategy is
that the fundamental structure of this state-space topology
changes depending on the exploiters choice of control parameter. This is a likely behaviour in real systems in which an
exploiter effectively combines a range of strategies dependent on current health of their system. The effect is to create a potentially uncertain and highly non-linear cascade of
resilience thresholds throughout the system, which must be
understood if an exploiter is to choose the optimal value of
their control parameter intelligently.
The exploiters choice of Sgoal is the primary management
strategy. Fig. 5(c) shows how the desirable steady state values
of the three economic indicators vary with the choice of Sgoal .
A value of Sgoal = 18.6 maximizes prot, while marginal prot
is maximized at maximum Sgoal .
The dynamic properties of the system are illustrated in
Fig. 5(d). Return time is minimized for Sgoal 14.0, while
resilience is maximized at minimum Sgoal . The plot of
resilience exhibits a jump discontinuity at Sgoal 16.9. In
general, there could be any number of discontinuities, of
any size, whenever a basin of attraction appears or disappears as a control parameter is gradually scanned across its
range. As an exploiters mental model and strategy becomes
more advanced, we might expect more and more structure
in the H isocline, and increasing potential for such threshold
effects. This highlights the importance of a dynamical systems model structure in the effort to understand feedbacks
in socialeconomicecological systems as a complement to
existing simulation and agent-based approaches. Misunderstanding, or absolute reliance on model systems that cannot reproduce threshold effects, may lead to unexpected and
catastrophic results from intuitive and linear management
strategies.
Fig. 5(e) illustrates the process that might cause an
exploiter to move their system across the threshold without
understanding the consequences of their actions. A grazier
currently sitting at point A may believe that a slight change
in strategy to point B might yield improved economic performance for little reduction in resilience. Obviously, however,
it is point C that they reach in their efforts to improve their

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Table 2 Dynamic properties when system managed at point of maximum prot

Strategy

Maximum prot (MP) (units H)

Cattle
Forage
Utilization
Compound

3.5
3.5
3.5
3.5

Residual N at MP (N)
50.3
50.3
50.3
50.3

system, with a potentially catastrophic reduction in resilience.

Note that this is not the same process as a system ip from
the desirable basin to the less desirable basin; it is more subtle.
The threshold passed is from a single-attractor, very resilient
system, to a dual attractor, less resilient system, with the
potential to undergo a catastrophic system ip. As the sudden transition does not affect the current value of a systems
state variables, it does not present any external indication that
it has occurred, making it particularly dangerous.

4.

Discussion

We have purposefully analysed four diverse exploiter strategies of the simplest feasible construction. Each system was
analysed as exploitation intensity was increased from the
unexploited case to a situation of maximum sustainable
intensity. In a constant environment, any of the operating
points we have investigated would be sustainable, and operating at the point of maximum prot or yield would be a rational
decision. In a real environment, which is variable and uncertain, the analysis of resilience across the sustainable management region summarizes the capacity of a system operating
at different levels of intensication and protability to persist.
There is a distinct tension between operating points that maximize prot and those that foster resilience in every system.
Table 2 shows that every strategy produces the same maximum long-term prot for the exploiter, with the same residual
stock of natural capital, because sustainable prot and production are fundamentally determined by the growth rate
of natural capital and the economic or biological parameters
describing how human-made capital processes natural capital to create production. Traditional economic analyses would
give a similar result in every case, not considering the impact
of the management strategy employed to drive the system
towards these long-term equilibria, therefore missing large
differences in dynamical behaviour away from equilibrium.
That the exploiters strategy changes the dynamic response of
the system to perturbations is well evidenced in Table 2 by the
resilience and return time for each strategy at the conguration generating maximum prot. The more detailed resilience
analyses illustrated that the opportunity cost of increasing
prot while reducing resilience varied strongly with management strategy.
Many modern and historical systems are characterized by
a strong feedback between prot and management strategy,
making the resilience of a system at the point of maximum
prot a particularly interesting quantity. It is important to note
that the resilience does not feed back to management strategy in the same way as prot because it is not an easily and
quickly measured quantity that can be used to inuence day-

Resilience at MP (N)
17.2
50.3
50.3
27.9

Return time at MP (years)

7.73
15.5
2.92
4.33

to-day management decisions. The only way to incorporate it

into real management decisions is by deliberately considering a model structure like the one we use here (Millennium
Ecosystem Assessment, 2005).
As a practical example, in exploitation systems exhibiting multiple steady states, MSY management can move the
system close to the edge of the desirable basin of attraction, drastically decreasing the resilience of the system. It
does maximize the sustainable yield of the system, but leaves
almost nothing for contingencies, a classic result from studies
of sheries (Clarke, 1990). In contrast, as the results show, in
a system with only a single basin of attraction, a signicant
amount of resilience to permanent qualitative change may
still remain when a system is managed at MSY. This might
indicate, for instance, that a strategy of constant utilization
would provide good resilience and good return time at optimum prot. However, we have also seen that if we model this
same system more realistically as a particular specication
of a compound management strategy, gradual tweaking of
management parameters in an effort to reach the point of
maximum prot can actually change the topological structure of state space. This implies that a system that has always
responded as if it contained a single basin of attraction might
invisibly cross a threshold to a multiple-basin state-space
topology. If a manager did not realize that they had passed this
threshold, it is possible that they would drive their system into
a non-desirable basin of attraction using only the successful
practices by which they had always run their system in the
past.
The resilience analysis calculated the maximum amount
of natural capital that could be removed from a system before
the management strategy drove a system to collapse. From
this perspective, a strategy that solely aims to maintain a
stock of human-made capital is unreactive, and strategies
that respond to the amount of natural capital in the system
are reactive. In the context of grazing systems, the rst scenario represents a constant stocking rate strategy, whereas an
exploiter that maintains a constant utilization rate manages
the ratio of human-made to natural capital and is perfectly
reactive. For a given system, a grazier who maintains a constant stocking rate despite a perturbation that removes half
their forage may drive the system to a collapse that would
be avoided by a grazier employing a reactive strategy. This is
reected by the very different resilience of the former strategy
compared to the latter strategies.
The analysis of the compound strategy allows us to generalize these results, and investigate how systems move from
one extreme to the other. It also represents a realistic general strategy within the context of family managed Australian
grazing systems, because: (1) stocking rate may be limited
below ecological carrying capacities due to labour constraints

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and to maintain quality of life, determining S2 ; and (2) almost

all stock may be sold off during periods of particularly bad
drought, determining S1 .
The analysis performed on the compound strategy investigated protability and resilience for different intensications
or stocking rates, S2 . The key result of this analysis was the
potential for such a system to exhibit a discontinuous threshold in resilience within the management region, to one side of
which resilience is maintained, while on the other resilience
is degraded. Whether this threshold exists within the accessible management region, or whether the system exhibits only
a high resilience or only a low-resilience state, is dependent
on the value of the remaining parameter, Sgoal .
Sgoal describes the reactivity of the exploiter, and is best
understood graphically. For given values of S1 and S2 , a reactive
exploiter will exhibit an almost linear H isocline across the
accessible region determined by environmental conditions,
max[N] K. In contrast, a relatively unreactive exploiter will
exhibit an H isocline with a distinct knee, as illustrated in
Fig. 5(a).
The resilience of such a system will be fostered by factors
that enhance the reactivity of the land manager, attening
the knee of the isocline such that it more closely approximates
the reactive linear isocline. The resilience will be degraded
by factors that encourage managers to maintain stock despite
perturbations that destroy natural capital. Fixed costs uncorrelated with stocking rate or natural conditions, such as interest repayments on traditional, inexible bank loans, will force
exploiters to keep stock on degraded land in an effort to maintain current output even at the cost of long-term land degradation.
Government subsidies to drought-affected grazing systems
present another example of economic effects that can modify system resilience. Importantly, different types of subsidies create very different outcomes. A subsidy that helps
drought-affected graziers purchase fodder to supplement
their degraded forage stocks encourages them to maintain
stocking rates despite effects that degrade natural capital
in the system. This exacerbates the knee in the H isocline,
potentially: (1) creating a resilience threshold; (2) moving an
existing threshold such that more of the accessible management region yields a low-resilience system; or (3) driving the
system into a regime in which only the low-resilience state
exists. Increased economic pressures during times of drought
will further compel a manager to drive the system towards
these regions of low resilience. Any further perturbation whilst
the system sits in this low-resilience conguration is increasingly likely to ip the system into an undesirable and lessproductive state, from which it will not recover even after
factors decreasing resilience are ameliorated.
An alternative subsidy is one that helps offset the transport costs associated with agisting cattle to distant ranches
unaffected by the drought. This encourages managers to react
to decreases in natural capital, attening the isocline. This
will tend to: (1) recover the possibility for the system to
exhibit a high-resilience state; (2) move the threshold such
that the high-resilience state occupies more of the management region; or (3) move the system into a regime in which
only the high-resilience state exists. This tactic helps ameliorate the plight of graziers during a time of drought, while

11

fostering resilience in the system so that even if they are

increasingly driven by the economic performance of their
system they are less likely to drive it into an undesirable
state.
Another change facing grazing systems is increasing
industrialization of what has previously being a hereditary exploitation system. Industrialization will yield different social drivers, removing, for instance, the emphasis on
lifestyle that currently limits exploitation intensity. This will
tend to increase the maximum stocking rate Sgoal towards values that maximize sustainable yield, perhaps also increasing
S2 in response to focus on short-term economic returns. Both
of these effects will tend to reduce the resilience of the system,
increase the likelihood of introducing thresholds of resilience
into the system, and increase the emphasis on economic performance likely to drive the systems across these thresholds.
These results apply equally well to other landscape
exploitation systems. A hunting-gathering society that limits
their population growth during plentiful times, through cultural and religious mechanisms, will be less likely to degrade
their systems during times of hardship (Anderies, 1998). Failing this, a society that responds to a decrease in available
natural capital, perhaps by splitting into smaller sub-groups
spread over a larger region, will be less likely to drive their system to collapse. In contrast, a society that generates a large
sunk-cost investment in administrative or religious buildings will be less likely to respond to environmental variation,
and more likely to drive their system to the edge of sustainable production (Janssen et al., 2003). For a system that has
reached this point, only a small economic, social or ecological perturbation may be necessary to push the system over
the threshold to collapse.
The long-term, undiscounted measures of prot and
resilience presented in the analysis represent one particular summary of system behaviour. Specically, they do not
illustrate the transient prot and resilience available to an
exploiter following a perturbation. This may be a critical factor if a perturbation temporarily affects an exploiters ability
to meet their xed costs, such that they must change their
strategy, emphasizing short-term prots and possibly further
degrading resilience at the time the system needs it most. Following a perturbation both instantaneous prot and resilience
will follow a trajectory back to their long-term steady state
values. Long-term protability and resilience are a summary
of this more complicated system behaviour, which provide
an important simplication allowing us to quantify at some
approximation the general response of the system.

5.

Conclusions

We have provided a quantitative analysis of the effects of management strategy on the non-equilibrium dynamical response
of generic systems harvesting renewable natural resources.
This is a powerful extension to previous works, providing
both solid numerical comparison, and an abstraction applicable across any system dependent on landscape exploitation.
This has allowed us to begin identifying fundamental social,
economic and ecological drivers affecting resilience in these
systems.

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Broadly speaking, management strategies responsive to

the stocks of a given state variable will foster the resilience
of a system to variability affecting that state variable. Mathematical verication of this intuitively logical statement can
be quickly invoked to estimate the dynamical costs and benets of various management strategies in real world systems,
from sunk cost effects in the early civilizations of Easter Island
(Brander and Taylor, 1998) and the Anasazi (Janssen et al.,
2003), to drought relief for modern graziers. Moreover, the
model allows the effect of alternative strategies to be calculated, and the opportunity cost of those strategies visualized
using multi-objective optimization.
This information is of vital importance if we are to manage the landscape exploitation systems that support our current civilizations (Millennium Ecosystem Assessment, 2005).
It is increasingly evident that the same social, economic and
ecological processes that drove collapse in civilizations such
as Easter Island are being repeated across the worlds modern exploitation systems, driving salinization, desertication
and landscape change (Brander and Taylor, 1998; Anderies et
al., 2002; Anderies, 2005; Millennium Ecosystem Assessment,
2005).
We will continue to make these same mistakes if we
try to independently manage each sector of the coupled
socialeconomicecological systems that drive progress in our
environmentally enlightened society, e.g., demanding sustainability or resilience at the same time as creating increasingly connected world-wide exploitation networks mediated
by free trade, without considering how these processes interact. We risk a particularly serious mistake when we apply
linear extrapolation to our conceptual, economic and mathematical models, failing to capture the possibility for threshold
effects including catastrophic system collapse.
Our model framework both shows and calculates how
protability and resilience trade-off against each other in
real systems, providing the information vital to choosing
how to exploit resources to support our societies. Of course,
Trading-off
maximum resilience cannot be the goal per se.
resilience for increased quality of life is a rational decision,
but we must at least attempt to be aware of the consequences
of the choices we are making when we do it (Millennium
Ecosystem Assessment, 2005). We need to improve our mental models, and this framework is an attempt to contribute to
that end.
The broad framework we have presented is a foundation:
we can more precisely capture every model sub-system when
studying specic systems. We already use more advanced
models of economic decision making based on discounted
utility over management time horizons. We are collecting data
on the social forces driving these decision making processes
within grazing systems in Australia, and more advanced ecological models of these systems already exist (Anderies et al.,
2002).
By allowing key parameters to evolve through time, or
become full state variables, it is possible to implement more
advanced descriptions of processes including technological
development (e.g., Reuveny and Decker, 2000; ca. Brander and
Taylor, 1998) and system adaptability (Walker et al., 2004). Initially, however, we plan to build on the strength of our very
simple models to inform system-wide descriptions being gen-

erated by multiple precise simulation models, and to help link

process to pattern in stochastic agent-based systems. The rst
synergy will combine the vital ability of our model to predict
thresholds and system-wide feedback effects with the precision of detailed but linear micro-scale models.
Most immediately, we are extending this local system
to an interacting spatial and networked mosaic to appropriately investigate resilience in regions heterogeneous in natural resources or controlled by variety of managers. This is
an essential component of regional resilience, where spatial
or strategic diversity might help foster the resilience of an
entire region to, say, climate change, while allowing individual
landowners to operate their systems for a reasonable prot.
The simple class of models investigated herein have
already identied three key behaviours of these systems: (1)
the choice of management strategy can drastically affect the
capability of a landscape exploitation system to respond to
variability or perturbations in the underlying social, economic
or ecological sub-systems; (2) within a given management
strategy, an exploiter will always have to make a subjective choice trading-off economic performance and resilience;
and (3) silent, unheralded changes in state-space topology
as a result of management decisions can leave a previously
resilient system balanced precariously close to disaster. Particularly disturbing is the potential interaction between the
last two effects in the modern environment of globalization
mediated by rapid, ultra-efcient economic communication,
unmatched by similar advances in social and ecological information transfer. A model like ours that provides information
about system-wide resilience, and the economic trade-offs it
requires, is the rst step to implementing balanced management strategies able to encompass system dynamics.

Acknowledgements
The authors thank Andrew Higgins, Peter Roebeling and John
Ludwig for input to the model and B. Walker for general discussions about resilience and adaptability. Nicky Grigg and Peter
Roebeling provided comments that improved an early draft of
the manuscript. This paper is a product of the project Identication of dynamically resilience exploitation-strategies in
complex landscape systems using spatially explicit dynamical
models and multi-objective optimisation funded by CSIROs
Complex Systems Science Centre.

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dynamics: the Tsembaga of New Guinea. J. Theor. Biol. 192 (4),
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