Total Environment Research Themes 3–4 (2022) 100011

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Total Environment Research Themes

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

Ecological dynamics, resilience and sustainability

William Grace
Australian Urban Design Research Centre, 1002 Hay Street, Perth, Western Australia 6000, Australia

A R T I C L E I N F O A B S T R A C T

Keywords: This article describes a simple generic system dynamics model of resource dependency in ecosystems to explore
Model the potential range of behaviours that can arise purely due to system structure. The model produces a small
Equilibrium number of behaviours that replicate the observed real‐world behaviour of ecosystems but does not produce
Ecosystem the less plausible outcomes produced in simpler predator–prey models (e.g. Lotka Volterra) such as exponential
Collapse
growth or exploding oscillations. The more limited range of behaviours is attributed to the inclusion in the
Predator
Prey
model of species dependence on abiotic resource flows. The simulations explore the response of systems in
Resilience equilibrium to various types of ‘shock’ and the outcomes are used to reflect on the concepts of resilience
Sustainability and sustainability. The results challenge the normative understanding of these terms and their relationship
to each other. The results also reinforce the notion that shocks to the abiotic system flows at the foundation
of all ecosystems present the greatest threat to ‘resilience’ as they affect all trophic levels.

1. Introduction may be value in informing research, the vagueness of the terms is

The concepts of sustainability and resilience share a common focus
‐ they both involve study of the dependency of agents on resources of
one type or another within a system, including in two of the well‐
known systems archetypes: limits to growth and tragedy of the com-
mons (Braun, 2002). Resource constraints are the reason ‘nothing
grows forever’. All animals, including humans, are dependent on a
wide range of biotic and abiotic resources. Predators depend on prey,
plants require air, water, minerals, and sunlight to grow.
The term ‘sustainability’ has in some cases been used interchange-
ably with ‘resilience’, a term credited to C.S. Holling from his seminal
article Resilience and Stability of Ecological Systems (C. S. Holling, 1973).
Although Holling’s early work was associated with understanding the
persistence of species in nature, the term ‘resilience’ has been
expanded by him and others since to include concepts such as ‘trans
formability’ and ‘adaptability’ in socio‐ecological systems (C.
Holling, Walker, Carpenter, & Kinzig, 2004; Carl et al., 2010). Several
articles have sought to draw connections between the terms ‘sustain-
ability’ and ‘resilience’ (Berkes & Folke, 1998; Fiksel, 2006; Ludwig,
Walker, & Holling, 1997; Redman, 2014), but none offer a satisfactory
explanation of the meaning of each term or their relationship to each
other in terms of system behaviour.
The lack of clarity about what is meant by these terms has led to
them being described as ‘boundary objects’, defined by some authors
as ‘a theoretical perspective explaining the role of objects in inter and trans-
disciplinary research’ (Lundgren, 2020). While it is accepted that there
unhelpful in promoting a general understanding of what is sustainable
and / or resilient and what is not. In order to answer this question, it is
necessary to have a model that reflects the observed dynamic beha-
viour of ecosystems in nature.
The biological resource dependency literature most often cited
relates to the interactions between predators and prey. A Google Scho-
lar search for ‘Predator prey models’ returns 900,000 results. Most of
these are mathematical treatments of the species interactions using dif-
ferential equations, the genesis of which is the widely cited Lotka‐
Volterra equations which are based on work originally published in
1910 (Lotka, 1910).
Swart (1990) reports on the insight gained from using system
dynamics modelling to explore system behaviour arising from the
Lotka Volterra equations. Swart’s model is configured as shown in
Fig. 1.
Models of this type produce a range of behaviours. Terminology
varies, but in this article the various terms have the following
meaning.
Equilibrium a condition in which stock (e.g. population) levels
are constant over time.
Oscillation periodic variation in stock levels which may or may
not vary in amplitude and / or frequency.
Periodic a form of oscillation in which the amplitude and
(continued on next page)

https://doi.org/10.1016/j.totert.2022.100011
Received 20 May 2022; Revised 30 August 2022; Accepted 30 August 2022

2772-8099/© 2022 The Author(s). Published by Elsevier B.V.

This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
W. Grace
orbit period is regular and repetitive.
Limit cycle a form of periodic orbit that is closed (i.e. “that a
point moving along the cycle will return to its
starting position at fixed time intervals and thus
execute periodic motion” Edelstein‐Keshet (1988)
Each of these conditions may be stable (i.e. the system has a tendency
to that condition independent of initial conditions and perturbations),
or unstable.
As reported by Swart, the Lotka Volterra structure can produce a
wide range of behaviours depending on its parameterisation. These
include exponential growth of both species, extinction of the preda-
tors, damped oscillations toward stable equilibrium, explosive oscilla-
tions, and non‐isolated closed orbits. Although the Lotka‐Volterra
equations are known for their ability to produce closed orbits, these
occur only with a specific set of parameters. As Swart notes: ‘In the neu-
trally stable Lotka‐Volterra model, both populations undergo constant oscil-
lations whose amplitudes bear no relation to the biology of the species but
only to the initial values.’ As identified by many other authors (Toro &
Aracil, 1988), the equations are insufficient to characterise any real
world examples of limit cycle behaviour, as the cycles created by the
equations are inherently unstable.
The existence of exponential growth of predators and prey is clearly
indicative of missing structure in the Lotka Volterra model. As nothing
can grow forever, the agents in question would be limited by balancing
feedbacks which are simply not represented in the model structure.
Similarly, exploding oscillations are rarely, if ever observed, again
probably because there are balancing feedbacks missing from the
model that would damp such behaviour.
Many others have produced mathematical models that modify and
expand the Lotka‐Volterra equations. The most well known variation
on Lotka Volterra is the Rosenzweig‐Macarthur model (Rosenzweig
& MacArthur, 1963). A version of that model after Turchin (1997) is
set out below in equation form.
 
dN N cNP
¼ r0N 1   ð1aÞ
dt K dþN
dP cNP
¼χ  δP ð1bÞ
dt dþN
where:
Fig. 1. Predator – prey model structure (Swart 1990) This simplified stock and flow diagram illustrates populations as ‘stocks’ and inflows / outflows from the
stocks as ‘rates of change’ in the stocks.
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Total Environment Research Themes 3–4 (2022) 100011
N and P are the populations of prey and predator respectively
r0 is the prey growth rate,
K is the prey carrying capacity,
c is the predator consumption rate
d is a constant regulating equilibrium (satiation) levels
δ is the predator death rate
χ is the predator–prey conversion rate
This configuration incorporates the so‐called “Holling Type‐II Func-
tional Response” which characterises how the predation rate responds
to prey density.
This model eliminates most of the obvious deficiencies in the pure
Lotka‐Volterra model and can produce a range of plausible behaviours
including: stable equilibrium populations of predators and prey,
damped oscillations leading to equilibrium, predator extinction and
limit cycles. In fact, the model produces two forms of stable limit cycle,
one in which populations oscillate in an elliptical manner similar to
the unstable Lotka‐Volterra orbits (Fig. 2a) and one in which limit
cycles ‘emerge’ with population levels that fall to almost zero (see
Fig. 2b). This behaviour occurs at elevated values of K – the prey car-
rying capacity, and is known as the ‘paradox of enrichment’
(Rosenzweig, 1971).
Apparently stable, limit cycle behaviour is observed in predator –
prey interactions in nature. Probably the most famous example of this
derives from fur records of lynx (predator) and snow hares (rey) in
Canada analysed by Elton and Nicholson(1942) based on data com-
piled by MacLulich (1937). The data is illustrated in Fig. 3 and illus-
trates approximately ten year cycles.
Other examples are reported in the literature including (Gilg,
Hanski, & Sittler, 2003) in respect of the collared lemming in the Arc-
tic tundra, and several others by Begon (2009) p.298. Replicating this
behaviour in the laboratory has been largely unsuccessful but recent
work by Blasius et al (2020) has produced limit cycles from interac-
tions between the aquatic invertebrate Brachionus calyciflorus (preda-
tor) and its prey, the green algal species Monoraphidium minutum
(prey). The evidence indicates that although stable limit cycles are
observed in nature, they cannot be reproduced by the simple Lotka‐
Volterra two‐species predator–prey model. As noted above, stable limit
cycles can be produced from the Rosenzweig‐Macarthur type models.
Charles Krebs from the Department of Zoology at the University of
British Columbia has published several papers on the lynx‐hare inter-
actions in conjunction with other authors. In (Stenseth, Falck,
W. Grace
Fig. 2.
Fig. 3. Hare-lynx limit cycle behaviour.
Bjørnstad, & Krebs, 1997), the authors report that a simple preda-
tor–prey interaction is insufficient to explain the observed behaviour.
“We have furthermore found an asymmetry in the way the lynx and the
hare are positioned within the ecosystem: The snowshoe hare appears to
be regulated from below and above (by a variety of Predators including
the lynx). The lynx, in contrast, seems to be regulated only from below,
and primarily by the hare. Thus, from the hare’s point of view, the food
chain is a vegetation–hare–Predator chain, whereas from the lynx point
of view, the hare–lynx interaction dominates.”
This suggests that any model that contains only predator–prey
interactions without prey‐food interactions may be inadequate to
reproduce all facets of species behaviour. Snow hares’ food resources
include a large variety of plant types which are also biotic resources
which in turn rely on photosynthesis, a process fuelled by abiotic
resources: air, water, sunlight, and nutrients (minerals). The availabil-
ity of air and sunlight can be considered as variable flows (at least at
temporal scales relevant to humans). Water and minerals cycle contin-
uously and so abiotic resources can be conceptually represented as
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Total Environment Research Themes 3–4 (2022) 100011
flows in an ecosystem model. In the Rosenzweig‐Macarthur model,
prey are regulated from below by the K value (see Equation 1) which
represents all lower trophic levels under the term ‘carrying capacity’.
Models commonly include the dynamic impact of such effects as “re-
source subsidies” (Leroux & Loreau, 2008; Nevai & Van Gorder,
2012) which may include allochthonous resources (e.g. carcasses that
seasonally float across the system boundary) or autochthonous
resources (e.g. dissolved organic matter in lakes (Lau, Sundh, Vrede,
Pickova, & Goedkoop, 2014)).
The term ‘resilience’, at least in respect of ecological systems, was
coined by C.S. (Buzz) Holling in his seminal (1973) article ‘Resilience
and Stability of Ecological Systems’. Holling was interested in the dif-
ferent forms of dynamic behaviour observable in the interactions
between species and their environment. In his terms, ‘stability’ refers
to ecological systems that are in a relatively stable dynamic equilib-
rium, meaning a return to equilibrium ‘after a temporary disturbance’.
By ‘resilience’ he meant ‘the persistence of systems and of their ability
to absorb change and disturbance and still maintain the same relationships
between populations or state variables’.
W. Grace
Fig. 4. Simplified stock and flow diagram of 2 level system.
The emergence of resilience theory has led to a significant research
community associated with the Resilience Alliance,1 spawning studies
spanning ecology, social and economic systems; and a dedicated journal
‐ Ecology & Society. Various modelling studies (Kar, Pal, & Ghosh, 2019;
Pal, Kar, Yamauchi, & Ghosh, 2020; Tromeur & Loeuille, 2017) have
considered the effect on resilience from human harvesting of resources,
most notably in marine ecosystems, where conventional management
policies pursue Maximum Sustainable Yield (MSY) of the target species
rather than resilience of the multiple species making up that ecosystem.
The objective of the work described here is twofold, firstly to create
a simple yet plausible generic model of ecosystems that explains
observed behaviour; and secondly to use this model to unpack the
meanings of, and relationship between, sustainability and resilience.
2. Methods
Here I describe a simple hierarchical system dynamics model
(Randers, 1997) of a food chain built sequentially in several phases to
explore the essential dynamics of biotic dependence on resources,
including the fundamental dependency of all life on abiotic resources.
The model is deliberately simplified in order to provide insights into
the generic behaviour of ecosystems based on the inherent structure of
dependency. The model has not been parameterised to reflect real world
behaviour of any specific species interactions, rather to explore the
range of possible system behaviours arising from that generic structure.
The model seeks to reflect the endogenous behaviour of such systems but
doesn’t include allochthonous resources or attempt to incorporate the
various autochthonous resources that exist in real systems.
The model was constructed using the Vensim Professional software
(Version 9.2.2 using a very small time step (0.03125) and R4K Auto
integration type to ensure model stability over a wide range of param-
eterisation. The model structure is set out below with further explana-
tion of the parameterisation of the model, in particular the stock
control functions, contained in Appendix A. The equations for the com-
plete model are included in Appendix B.
2.1. Biotic hierarchy modelling
2.1.1. Model construction
In order to approximately replicate the notional sequential intro-
duction of biotic resources in a landscape (i.e. plants followed by her-
bivores followed by predators), the model was constructed in discrete
steps.
1
https://www.resalliance.org/.
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Total Environment Research Themes 3–4 (2022) 100011
The initial stage model simulates plant dependency on soil mois-
ture, and parameters were varied (including with extreme values) to
test system behaviour, namely: the relative growth rates of plants
and soil moisture, the unit consumption rate of moisture by the plants
and the graphical functions that regulate the inflows and outflows to
the stocks. The parameters were then finally fixed at the parameter
set that produced a smooth dynamic transition to equilibrium. The
model was then extended to add herbivores at a time after equilibrium
had been established between plants and soil moisture. Parameters
relating to the behaviour of herbivores were then varied to test system
behaviour patterns, as in the previous step. Once again, parameters
were then finally fixed at the values that produced a smooth transition
to equilibrium across the expanded system. Thereafter, predators were
added and the process repeated. Specific details of each step are set out
in the following sections, and the equilibrium parameter values are
included in Appendix B.
2.1.1. Plant dependency on Soil Moisture
Consider a barren field that is colonised by a single plant species (a
biotic resource) that is dependent (inter alia) on Soil Moisture (an abi-
otic resource).
Fig. 4 depicts the dynamics of this two‐level system, which can be
read as follows:
Loop B1 The stock of Soil Moisture (SM) is (on average)
constant over time prior to plant colonisation, i.e. there
is zero inflow. As SM is consumed by plants it is
replenished by inflows regulated by function fSR (SM)
up to a maximum rate (RSM).
Loop B2 SM is depleted by moisture transpired by the Plant
Mass (PM) at a rate controlled by its maximum
transpiration rate (CSM), regulated by function fSC(SM).
Loop B3 The Plant Mass (PM) regeneration rate is a function
fPR(SM) of Soil Moisture up to a maximum regeneration
fraction (RPM) and regulated by Loop R1.
Loop R1 The product of the regeneration rate and the stock
value of PM.
dSM
¼ RSM  f SR ðSM Þ  PM  CSM  f SC ðSM Þ ð2Þ
dt
dPM
¼ PM  RPM  f PR ðSM Þ ð3Þ
dt
W. Grace Total Environment Research Themes 3–4 (2022) 100011

where. In this configuration the equation for Plant mass is modified as

shown in Equation (6) and the equation for Herbivores is added (Equa-
RSM, CSM, RPM are non‐zero constants tion (7)).
fSR is a decreasing function with domain {0 ≤ SM/SMi ≤ 1} and
range {0 ≤ fSR ≤ 1} dPM
¼ PM  RPM  f PR ðSM Þ  H:CPM :f PC ðPM Þ ð6Þ
fSC is an increasing linear function with domain {0 ≤ SM/SMi ≤ 1} dt
and range {0 ≤ fSC ≤ 1}
fPR is an increasing linear function with domain {0 ≤ SM/SMi ≤ 1} dH
and range {‐1 ≤ fPR ≤ 1} ¼ H:RH :f HR ðPM Þ ð7Þ
dt

Images of the functions are included in Appendix A, and explained At equilibrium.

further in the following.
RPM :f PR ðSM Þ
fSR regulates the inflow to Soil Moisture in response to its stock H ¼ PM: ð8Þ
level. At t = 0 soil is saturated, and therefore fSR = 0, meaning there CPM :f PC ðSM Þ
is no inflow, thus the system starts in equilibrium. As Soil Moisture is The additional functions fPC (that regulates Plant Mass outflow in
depleted by plants through evapotranspiration, Soil Moisture is replen- response to consumption by Herbivores) and fHR (that regulates Herbi-
ished and fSR > 0, meaning there is a non‐zero inflow whenever Soil vore inflow in response to Plant Mass) are illustrated graphically in
Moisture falls below its initial value. Appendix A. fPC is identical to fSC and fHR is structured similarly to
fSC regulates the outflow from Soil Moisture. Initially there is no fPR. CPM and RH are constants.
restriction to outflow (i.e. fSC = 1) but as Soil Moisture is depleted,
the outflow from the stock diminishes. Should Soil Moisture be com-
pletely depleted, fSC = 0, the outflow is zero (which ensures that Soil 2.1.3. Predator dependency on Herbivores
Moisture stock cannot fall below zero). Consider now a further resource dependent agent, a Predator,
fPR regulates the inflow to the Plant Mass stock in response to the entering the system (Fig. 6) after equilibrium has been reached
Soil Moisture level. This inflow represents the net regeneration rate between Herbivores, Plants and Soil.
of Plant Mass and therefore must have a positive and negative range The equations for this system follow.
(set here to −1 to + 1) in order for the stock of Plant Mass to both
increase and decrease. The inflow must be zero at equilibrium (i.e. dH
¼ H  RH  f HR ðPM Þ  Pr:CPr :f HC ðH Þ ð9Þ
when fPR = 0). As configured here, this occurs when Soil Moisture dt
reaches a quarter of its initial value.
At equilibrium: dPr
¼ Pr:RPR :f PR ðH Þ ð10Þ
dt
RSM  f SR ðSM Þ
PM ðt Þ ¼ ð4Þ
CSM  f SC ðSM Þ And at equilibrium.

RH :f HR ðPM Þ
f PR ðSM Þ ¼ 0 ð5Þ PR ¼ H: ð11Þ
CPR :f HC ðH Þ
The additional functions fHC (that regulates Herbivore outflow in
2.1.2. Herbivore dependency on plants response to predation) and fPr (that regulates Predator inflow in
Consider now the situation where plants have been established and response to Herbivore stocks) are illustrated graphically in Appendix
the system is in equilibrium, when a small number of Herbivores enter A. fHC is identical to fSC and fPC, and fPr is structured similarly to fPR
the area. This expands the model structure as illustrated in Fig. 5. and fHR. CPr and RPR are constants.

Fig. 5. Simplified stock and flow diagram of 3 level system.

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W. Grace
Fig. 6. Simplified stock and flow diagram of 4 level system.
2.2. Resilience testing - post-equilibrium disturbances
To explore the concept of resilience requires modelling of the sys-
tem response to disturbances or shock. This was tested by introducing
changes after equilibrium has been achieved in the hierarchy depicted
in Fig. 6.
3. Results
3.1. Plant and Soil Moisture interaction
The two stock model described in Fig. 4 starts in equilibrium with
zero net inflow and outflow of soil moisture (i.e. the soil is saturated).
Plant mass is introduced at time zero with an initial small stock. This
commences the dynamic responses of Plant mass and Soil moisture.
Sensitivity testing identifies that the behaviour of this system configu-
ration is limited to a small number of outcomes.
When unconstrained by limitations in Soil Moisture the Plant Mass
expands at its maximum growth rate (RPM). Eventually the growth rate
of the Plant Mass becomes constrained by limitations in Soil Moisture,
i.e. the Plant Mass regeneration rate drops to zero. At this stage Plant
Mass and Soil Moisture are in equilibrium, meaning the rates of inflow
and outflow of Soil Moisture are equal. All of this assumes the only
impact on the stock of Plant Mass is Soil Moisture.
The system’s eventual equilibrium is reached when fPR = 0 (Equa-
tion (4)). As parameterised, at that point fSC = 0.25 and fSR = 1 which
means that Soil Moisture (SM) is a quarter of its initial value and
inflow and outflow are identical at the rate of RSM. Accordingly, the
product of Plant Mass (PM), fSC and CSM is equal to RSM. Increasing
or decreasing RSM changes the equilibrium value of the Plant Mass
(PM) but otherwise has no effect.
The dynamic behaviour of the system is controlled by the relative
magnitudes of RPM (the growth fraction of Plant Mass) and RSM (Soil
Moisture inflow rate) ‐see Fig. 7. Below a certain ratio of those param-
eters (RPM / RSM) the system exhibits a smooth transition to equilib-
rium, irrespective of other system parameters. However, above that
threshold the system exhibits damped oscillations prior to equilibrium
becoming established at the same stock values.
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Total Environment Research Themes 3–4 (2022) 100011
At very high ratios of RPM to RSM, the amplitude of the oscillations
becomes so extreme that Plant Mass is diminished almost to zero, i.e.
overshoot and collapse behaviour is apparent (Fig. 8). However, as
noted above, Soil Moisture cannot be indefinitely depleted, and so this
apparent behaviour is misleading. If the timeframe of the model is
extended, it can be seen that what appears to be overshoot and col-
lapse in Fig. 8 is actually very long period damped oscillations leading
eventually to equilibrium at the same stock levels of Plant Mass.2
Examination of Equation (3) illustrates that an equilibrium level of Plant
Mass (PM) cannot fall to zero indefinitely. As Plant Mass grows, Soil
Moisture is depleted but is continually replenished, even if the stock
level reduces to zero. This is the outcome that results from the inflow
to Soil Moisture being a ‘rate’ rather than a fraction of the stock level.
In summary, the Plant Mass ‐ Soil Moisture system inevitably trends
to a unique equilibrium, albeit with damped oscillatory behaviour evi-
dent with some combinations of variables. Extreme value testing of the
model parameters confirms that other modes of behaviour such as
exponential growth, limit cycles or exploding oscillations cannot be
produced with this model structure.
This behaviour is reflective of the dependent resource being an abi-
otic rather than biotic resource. The inflow (growth rate) of a biotic
resource is some fraction of the stock level (i.e. the higher the stock
level the faster the growth rate and vice versa) which creates reinforc-
ing feedback. If the Plant Mass was to fall to zero (e.g. through a catas-
trophic fire) the Plant Mass would become locally extinct. In contrast,
the inflow to an abiotic resource reduces with the stock level (i.e. the
higher the stock of Soil Moisture the slower the rate of inflow) creating
balancing feedback. If the Soil Moisture were to fall to zero (e.g. years
of drought) it would recover as soon as rainfall returned. This simple
model exemplifies the dependence of all photosynthetic organisms
on the abiotic resources of sunlight, nutrients and water.
2
Of course in the real world this level of depletion may lead to extinction of Plant mass,
the atto‐Fox problem.
W. Grace
Fig. 7. 2 level system - dynamic behaviour arising from variation of RPM to
RSM.
Fig. 8. 2 level system - overshoot and collapse arising from extreme ratio of
RPM to RSM.
3.2. Herbivore, Plant and Soil Moisture interaction
Herbivores are introduced to the model at time 50 (the three stock
system described in Fig. 5) with the Plant mass and Soil moisture
parameters set at the values that produced a smooth transition to equi-
librium in the previous stage (see Fig. 7).
The behaviour of is dominated by the response of Herbivores to the
availability of Plant Mass. This is regulated by the Herbivore regener-
ation fraction (RH. fHR), or more specifically the level of Plant Mass that
can sustain a positive Herbivore growth rate. If the Plant Mass is insuf-
ficient to create a positive growth rate (births minus deaths) the Her-
bivores immediately decline to extinction (zero population). Above
that threshold value, the system comes to equilibrium smoothly
(Fig. 9) or via damped oscillations, depending on the relative values
of RH and RPM (Fig. 10). The higher this ratio, the more the system
overshoots as it seeks equilibrium. Nevertheless, the oscillations are
damped.
However, if the level of Plant Mass that can sustain a positive Her-
bivore growth rate is lower than the threshold level, the system
reaches a bifurcation point and exhibits overshoot and collapse beha-
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Total Environment Research Themes 3–4 (2022) 100011
Fig. 9. 3 level system – smooth transtion to equilibrium.
Fig. 10. 3 level system – oscillations arising from increasing ratio of RH to RPM.
viour as the plant resource is exhausted (Fig. 11). These outcomes
result from the parameterisation of the graphical function fHR, more
specifically the Plant Mass level that triggers a negative Herbivore
growth rate (i.e. population decline). The difference in Herbivore
response to Plant mass scarcity between these three situations, i.e.
immediate decline, equilibrium or overshoot and collapse, is depicted
in the function fHR for each case (Fig. 12).
If a (relatively) fast Herbivore growth rate is combined with a
‘slow’ response to resource scarcity then overshoot and collapse beha-
viour transitions to very long period and deep stable limit cycles
(Figs. 13a and b). These limit cycles are of the form evident in the
phase plot shown in Fig. 2b).
This behaviour (Fig. 13) occurs at the higher (relative) Herbivore
growth rates because the collapse happens so quickly that the Herbi-
vore stock declines sufficiently to be negligible before the Plant Mass
is completely exhausted, leading to a slow recovery of Plant Mass
and concomitant (eventual) recovery of Herbivores. In the overshoot
and collapse case (Fig. 11), the Plant Mass is exhausted first, condemn-
ing Herbivores to extinction.
Extreme value testing of the model parameters illustrates that this
structure does not produce exponential growth or exploding oscilla-
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Fig. 11. 3 level system – overshoot and collapse arising from extreme ratios of
RH to RPM.
Fig. 12. 3 level system – graphical function fHR(PM).
Fig. 13a. 3 level system – limit cycles arising from extreme ratios of RH to RPM
combined with fHR(PM) = 0 at lower levels of PM.
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Total Environment Research Themes 3–4 (2022) 100011
Fig. 13b. 3 level system – phase plot associated with Fig. 13a).
tions, nor stable limit cycles of the Rosenzweig‐Macarthur model type
illustrated in Fig. 2a).
3.2. Predator, Herbivore, plant and Soil Moisture interaction
Predators are introduced to the model (the four stock system
depicted in Fig. 6) at time 200 with the Plant mass, Soil moisture
and Herbivore parameters set at the values that produced a smooth
transition to equilibrium in the previous stage (see Fig. 9).
The behaviour mirrors the earlier introduction of Herbivores, i.e. it
is dominated by the response of Predators to scarcity of Herbivores.
Above the threshold that leads to equilibrium, the system either does
so smoothly (Fig. 14) or with damped oscillations (Fig. 15). As for the
previous Herbivore / Plant mass case, below that threshold, the system
exhibits either: overshoot and collapse (Fig. 16) of both Predators and
Herbivores; or very long period deep limit cycles (Fig. 17). Again,
extreme value testing confirms that neither exponential growth or
exploding oscillations are evident with this parameterisation of the
model, nor stable elliptical limit cycles. As for the previous case only
very long period and deep stable limit cycles are produced by the
model.
Fig. 14. 4 level system – smooth transition to equilibrium.
W. Grace
Fig. 15. 4 level system – oscillations arising from increasing ratio of RPr to RH.
Fig. 16. 4 level system – overshoot and collapse arising from extreme ratios of
RPr to RH.
Fig. 17. 4 level system – limit cycles arising from extreme ratios of RPr to RH
combined with fPr(H) = 0 at lower levels of H.
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Total Environment Research Themes 3–4 (2022) 100011
3.3. Resilience testing
Persistence of the system in stable equilibrium (as shown in Fig. 14)
was tested by introducing a ‘spike’ in each stock of around 20 % of its
equilibrium level at time 600. This level of disturbance has no long‐
term impact on the system, whether or not the spike is an increase
or decrease. In each case stock levels return to their initial equilibrium
levels with minor damped oscillation. Fig. 18 represents a sudden
increase in Herbivore numbers, which initially triggers increases in
Predators and decrease of Plant Mass. This confirms that the equilib-
rium that is established is stable.
The next test of stability is the response of the system to slower
changes in abiotic flows. This case was tested by slowly increasing
and decreasing (through ramping) the flows that regulate Soil Mois-
ture, following the establishment of equilibrium. Decreasing the inflow
of Soil Moisture has the predictable effect of reducing Plant Mass and
therefore both Predator and Herbivore numbers, with Predators at the
top of the trophic scale being most impacted (Fig. 19). At larger reduc-
tions of Soil Moisture flows, Predators become extinct (Fig. 20) and at
even larger reductions so do Herbivores. This begs the question about
whether resilience is a property of the system or the species. The land-
scape has returned to its original form, but animals have vanished.
4. Discussion
The simulations demonstrate that system behaviour is determined
by:
• The relative maximum (i.e. unconstrained) population growth rates
of the dependent agent and its resource, in combination with:
• the dependent agent’s response to extreme resource scarcity, i.e.
the resource level at which the agent’s population growth rate
becomes negative through increased mortality and / or reduced fer-
tility – referred to here as ‘growth rate zero’.
There are distinct categories of agent behaviour observed in the
simulations (see Fig. 12 and Table 1).
The deep limit cycles evident in the experiments where agent and
resource populations decline to near zero, occur near the ‘growth rate
zero’ boundary between a transition to equilibrium and overshoot and
collapse, and are influenced by the unconstrained growth rate of the
dependent agent.
For systems exhibiting immediate decline or a transition to equilib-
rium, the unconstrained growth rates of the dependent agent affect the
Fig. 18. Post equilibrium Herbivore injection.
W. Grace
Fig. 19. Gradual small reduction in Soil Moisture.
Fig. 20. Gradual larger reduction in Soil Moisture.
rates of population growth/decline and oscillatory behaviour (which
occurs at higher relative growth rates) but not the equilibrium magni-
tude of the stocks.
I have referred to the observed limit cycles as ‘deep’ as they are not
(stable) Rosenzweig‐Macarthur type elliptical cycles in which popula-
tion levels remain well above zero (Fig. 21), or oscillations that arise
Table 1
Summary of simulation results.
Growth rate zero
If ‘growth rate zero’ occurs above a resource level which is insufficient to
support a positive growth rate from the outset of the agent’s emergence
If ‘growth rate zero’ occurs below a resource level at which continuing agent
population growth leads to severe depletion of the resource stock.
If ‘growth rate zero’ occurs at a resource level between the upper threshold and
a lower threshold.
Total Environment Research Themes 3–4 (2022) 100011
Fig. 21. Phase plot of stable elliptical cycles (modelled).
due to cyclical variation in resource availability. In these deep cycles,
agents and resource stocks decline to near zero (Fig. 22).
Continuous models (such as that described here) that can produce
very small population levels are sometimes referred to as having the
“atto‐fox problem”, a term that derives from criticism by Mollison
(1991) of a model by Murray and Seward (1992) of fox rabies in which
foxes decline to 10−18 (atto‐) foxes per square kilometre. In the real
world such deep cycles will in many cases likely lead to extinction.
Nevertheless, these deep limit cycles seem to be consistent with the
cycles identified in the Hare‐lynx data (see Fig. 23), the Blasius
(2020) data (see Fig. 24) and those described by Holling in the spruce
budworm example.
These occur when the decline of the dependent agent in response to
critical resource scarcity is so fast that population levels approach zero
before the resource stock is completely depleted (compare Figs. 11 and
13a). Accordingly, both stocks eventually recover and the cycle is
repeated. These cycles are also referred to as ‘relaxation oscillations’
that occur when the growth rates of the dependent agent and resource
vary significantly in so called slow‐fast systems (Saha, Pal, & Banerjee,
2021). In contrast, when the response to scarcity is ‘slower’, the
resource stocks decline to zero first, leading to overshoot and collapse,
ensuring the extinction of both species (see Figs. 11 and 16).
Rosenzweig‐Macarthur type models can produce a similar type of limit
cycle but in that case it is associated with the resources available to
prey at high levels of carrying capacity (“nutrient enrichments”) rather
than their response to predation.
Relative unconstrained System behaviour Descriptor
population growth rates
Influences rate of decline Exponential decay of initial dependent agent Immediate
only population to zero decline
Low Dependent agent and resource populations Overshoot
decline to zero and collapse
High Dependent agent and resource decline to near- Deep limit
zero and slowly recover cycles
Low Smooth transition to a stable equilibrium for Transition to
dependent agent and resource populations Equilibrium
High Damped oscillations to a stable equilibrium for
dependent agent and resources populations
10
W. Grace
Fig. 22. Phase plot of deep limit cycles (modelled).
Fig. 23. Phase plot of Hare Lynx data cycles.
As noted above, the empirical data seems to support the existence
of deep limit cycles (relaxation oscillations) but it is interesting to
question whether there is empirical evidence of stable elliptical limit
cycles such as those produced by Rosenzweig‐Macarthur type models.
The model presented here does not produce oscillations of that kind,
nor have they been identified in the literature reviewed for this study.
All real‐world systems mimic the essential hierarchical resource
dependency described in these simple experiments, although with
much more complexity, including competition between species at each
level, omnivores operating at both the Herbivore and Predator level,
and with fungi, bacteria, insects and birds affecting all levels together
with irregular allochthonous or autochthonous resource subsidies.
Despite the simplifications, the experiments replicate the real‐world
behaviour observed in nature including: smooth transitions to equilib-
rium, damped oscillations, overshoot and collapse, and deep limit
cycles and (Begon, 2009; C. S. Holling, 1973; Klein, 1968). Exploding
11
Total Environment Research Themes 3–4 (2022) 100011
Fig. 24. Phase plot of Blasius data cycles Data from the C2 experiments.
oscillations or unconstrained exponential growth which are found in
the original Lotka Voltera model are neither evident in nature or in
these experiments. Other more sophisticated models derived from
Lotka Volterra eliminate those problems but retain some behaviours
that are not reproduced in the complete hierarchical model presented
here (e.g. stable elliptical limit cycles).
It is posited here that the reason for plausible behaviour is the
inclusion of abiotic flows (in this case water) at the base of all ecosys-
tems. This means the regeneration of this abiotic resource is regulated
by a balancing feedback loop, whereas in most predator–prey models,
regeneration of the prey is the product of the stock itself and its growth
fraction, which is a reinforcing feedback loop. This suggests that the
modelling of biotic systems without their underlying dependency on
abiotic resources such as air, water, nutrients (minerals) and solar radi-
ation may be insufficient to explain observable behaviour in the field.
This is not to say that other models could not be configured in a way
that gives rise to similar outcomes.
Studies on resilience such as those conducted in fisheries manage-
ment (Pal et al., 2020) might benefit from a more explicit inclusion of
species hierarchy such as in the model presented here.
5. Model validation
The process of model validation is to build confidence in its ability
to achieve its particular purpose (Schwaninger and Groesser, 2020;
Senge & Forrester, 1980). The model presented here can be thought
of as in ‘insight’ model, or a ‘causal‐descriptive’ model (Barlas,
1996). Unlike many system dynamics models it is not aimed at identi-
fying the “effectiveness of alternative policies”; its purpose is to
explore the set of behaviours that emerge from a plausible generic
model structure. It does not seek to replicate the exact dynamic beha-
viour of a specific hierarchical food chain, rather to depict the condi-
tions that give rise to the persistence or otherwise of each level in the
chain. Accordingly, it has not been calibrated against empirical data
for any particular set of species, nor contains additional structure to
represent the myriad of interdependencies that exist in real ecological
systems, most obviously competition at each trophic level. However,
the general behaviour produced by the model can be tested against
W. Grace
the empirical evidence, including in respect of predator‐ prey dynam-
ics and the existence of limit cycles. Establishing its consistency with
that evidence is the prime purpose of the study, and citations to the rel-
evant literature are included in the text of the article to support its
achievement of that aim. Because the model does not seek to simulate
specific real‐world species interactions, questions of boundary ade-
quacy (i.e. does the model contain sufficient structure to satisfy its pur-
pose) are primarily satisfied by producing plausible behaviours, i.e.
those observable in nature.
The key to the model behaviour is the parameterisation of the
inflows and outflows to the stocks, structured as constants modified
dynamically through graphical functions. With the sequential addition
of each level of the hierarchy, the parameters and graphical functions
were iteratively varied to evaluate the “window” within which the
dependent agent could establish, and extreme value testing used to
explore the full range of system behaviours following establishment
– effectively representing behaviour sensitivity testing at each stage
of model development. The model does not purport to replicate the
actual growth rates of the individual stocks, rather it establishes the
relative rates between each agent that govern persistence or otherwise.
Thus, the development of the model incorporated both parameter ver-
ification and extreme value testing, while being dimensionally
consistent.
6. Conclusions
A simple model of a four‐level ecosystem provides insights into
plausible system behaviour (i.e. that which is observable in the real
world). It is argued here that this is due to the founding of the biotic
resources on an abiotic flow, in this case water, which creates a balanc-
ing feedback loop at the lowest system level and limits the behaviour
of higher trophic levels to a small number of outcomes, namely equi-
librium achieved in a smooth transition or with tightly damped oscil-
lations, overshoot behaviour leading to collapse, or deep limit cycles.
As all life on earth is founded on such flows, this suggests that any
ecosystem models that neglect such flows may in some cases produce
unrealistic system behaviour.
Second, that stable limit cycle behaviour can be produced by sys-
tem structure alone, in the circumstances where overshoot does not
quite create collapse, but rather leads to very small stock levels
and slow recovery. These cycles may initially expand in amplitude
but inevitably settle into stable limit cycles. Holling’s original 1973
paper suggested that the budworm / balsam fir interactions were
an example of ‘…a distinct domain of attraction determined by the inter-
action between budworm and its associated natural enemies, which is peri-
odically exceeded through the chance consequence of climatic conditions’.
While that interpretation is not disproven by these experiments, they
do illustrate that this behaviour can be created by the same system
structure that produces other observable behaviour patterns without
random variations.
Dynamic equilibrium conditions are observable in nature, and so
are the damped oscillations that occur in response to natural varia-
tion and disturbances. Are such systems therefore ‘sustainable’? It
is more helpful to consider whether the outflows from dependent
stocks are sustainable, i.e. whether net consumption of the dependent
agent’s resource can be sustained over time. This depends both on
the net depletion rate of the resource (i.e. population levels and con-
sumption rates of dependent agents), and the stability of the inflows
(regeneration rates) to all the resources at lower trophic levels,
including the abiotic resources that support all life on earth. Accord-
ingly, sustainability is a time limited concept. Nothing stays the same
forever, climate conditions change, systems re‐configure on the
appearance of new agents, species become extinct through disease,
and so on.
12
Total Environment Research Themes 3–4 (2022) 100011
Holling defined resilience as ‘..the persistence of systems and of their
ability to absorb change and disturbance and still maintain the same rela-
tionships between populations or state variables’.
This definition implies a system in dynamic equilibrium prior to a
disturbance; in the case of the budworm ‘the chance consequence of cli-
matic conditions’. However, as these experiments show, the ability of
a species within an ecosystem to ‘persist’ is merely a function of the
scale and nature of the disturbance. If the disturbance is to the fun-
damental abiotic flows or lower trophic levels, the impact on the
higher levels is larger. Such disturbances may lead to the extinction
of one or more species in an ecosystem while leaving others in place,
albeit in a reconfigured equilibrium. Such a change would not meet
Holling’s resilience test as the ‘same relationships between populations’
would no longer exist. However, later papers by resilience theorists
have proposed a difference between ‘specific resilience’ and ‘general
resilience’. In somewhat of a summary of resilience and related con-
cepts Folke et al (2010) describe the former as ‘resilience of some par-
ticular part of a system, related to a particular control variable, to one or
more identified kinds of shocks’ and the latter as ‘the resilience of any
and all parts of a system to all kinds of shocks, including novel ones.’ This
represents somewhat of a qualification of the original Holling
definition.
The results presented here also question whether resilience is a
property of a ‘system’ or a specific species within a system. It could
be said that (for example) plants are resilient to virtually any shock
(prior to the appearance of Herbivores) if they are underpinned by a
constant flow of Soil Moisture. However, the resilience of the animals
at a higher trophic level is more limited.
The focus of the resilience community is on human intervention
intended to retain the persistence of a given system (including so
called socio‐ecological systems3). Based on the findings set out here,
intervening in systems subject to some specific shock (i.e. specific resili-
ence) would seem to be the more fruitful endeavour. As noted elsewhere
(Willliam Grace & Pope, 2015) no system is likely to withstand a direct
asteroid hit or catastrophic climate change. Therefore, what is referred
to as general resilience is really just specific resilience to a wide range
of plausible disturbances.
It is over thirty years since the Brundtland report “Our Common
Future” (World Commission on Environment and Development,
1987) produced the oft‐quoted explanation of sustainable develop-
ment: 'development that meets the needs of the present without compro-
mising the ability of future generations to meet their own needs'. If our
prime interest is the ‘persistence’ of humanity with an aspiration to
meet the Brundtland objective (i.e. the achievement of a decent stan-
dard of living for all), then the global socio‐ecological system must
first achieve a stable dynamic equilibrium. Exponential increases in
population and the increasingly unsustainable depletion of renewable
and non‐renewable resources indicates that humanity is nowhere
near such an equilibrium, as illustrated by the Limits to Growth stud-
ies first published in 1972 (Meadows, Meadows, Randers, & Behrens,
1972) and is now in a state of overshoot as more recent work
explains (William Grace, 2015; Meadows, Meadows, & Randers,
2004). Sustainability can be thought of as a quest for such an equi-
librium. Resilience of the human (or any) species can be thought
of as the ability to recover from shocks and disturbances once equi-
librium has been achieved.
Shocks and disturbances to the lower trophic levels will have more
impact on humans (e.g. the loss of pollination services (Kevan & Viana,
2003)), than those to higher trophic levels. However, shocks to the
abiotic resources that underpin life on earth will have even more
impact, as they threaten the entire dependent biotic system. Changing
3
This descriptor is open to the accusation of tautology as humans are obviously part of
ecological systems.
W. Grace Total Environment Research Themes 3–4 (2022) 100011

the radiation balance in the earth’s atmosphere is just such a distur- Appendix B. Model equations
bance, making climate change our most urgent challenge, albeit not
the only one. The model behaviour is dependent on the graphical functions and
constants, which were set to ensure the biotic agents (Plant Mass, Her-
Declaration of Competing Interest bivores and Predators) could reach a stable equilibrium after their
sequential appearance in the model.
The authors declare that they have no known competing financial The key variables that affect behaviour are the constants that con-
interests or personal relationships that could have appeared to influ- trol stock inflow (regeneration) rates, i.e. RSM, RPM, RH, and RPR. A mul-
ence the work reported in this paper. tiplier for each of these constants was included in the model to
facilitate sensitivity and extreme value testing of the model. The
Appendix A. - model parameterisation results of these tests are described in the article text.
The following sets out the equations for the full 4 level model. This
Graphical functions. should be read in conjunction of the figures and equations in the text.
Equations are set out in the order that the model was constructed, i.e.
fSR = GF1 (SM/SMi)
With consistent rainfall and without plants, Soil Moisture will • Plant mass and soil moisture interactions
vary through the year but reach an equilibrium state on an • Additional interactions following introduction of Herbivores
average annual basis. It is assumed in this model for simplicity • Additional interactions following introduction of Predators
that this is saturation (fSR = 0) so the model commences in
equilibrium. As Soil Moisture declines below a threshold, inflow The full Vensim model is available on request.
increases to a maximum value of RSM. Modelled as a linear
function but testing with non‐linearity show minor impact on Model settings
behaviour
fSC = GF2 (SM / SMi)
Conventional stock control. Soil moisture outflow is linearly
constrained as stock level falls. Non‐linearity affects the INITIAL TIME = 0
equilibrium level of Plant Mass. This is because concave GF2 Units: Year
constrains Soil Moisture loss – keeping Soil Moisture stock The initial time for the simulation.
higher, and hence plant regen higher for longer. Convex GF2 FINAL TIME = 200
causes the opposite. Essential model behaviour is though Units: Year
unchanged. The final time for the simulation.
fPR = GF3 (SM / SMi) TIME STEP = 0.03125
Plant mass regeneration rate is constrained once SM / SMi falls Units: Year [0,?]
below 0.5. Equilibrium obtained when fSR = 0 at SM / The time step for the simulation.
SMi = 0.25. Regeneration rate becomes negative at lower levels
of SM / SMi. Effect of non‐linearity: convex GF3 slows the
transition to equilibrium as restriction on PM inflow occurs
earlier. Concave is the opposite. Essential model behaviour is Soil moisture equations
though unchanged.
fPC = GF4 (PM / PMi)
As for GF2
fHR = GF5 (PM / PMi)The domain determines the envelope of Soil moisture = INTEG (SM inflow‐SM outflow, Initial soil
persistence of Herbivores. If there is insufficient Plant Mass to moisture)
support a positive Herbivore generation rate then the Herbivore Units: SM perc
influx immediately declines to zero. At the other extreme, if the Initial soil moisture = 200000
transition from positive to negative regeneration rates occurs at Units: SM perc
a lower bound threshold, overshoot and collapse SM inflow= “Max SM inflow (Rsm)”*”fSR(SM)”*SM Multiplier
occurs.As for GF3, non‐linearity has no significant impact on Units: SM perc/Year
essential behaviour (assuming zero point is maintained) “Max SM inflow (Rsm)”= 50,000
. Units: SM perc/Year
fHC = GF6 (H / Hi) SM Multiplier = 1
As for GF2 and GF4 Units: Dmnl
FPr = GF7 (H / Hi)As for GF5. If the transition from positive to [Note this variable was used to vary Rsm for sensitivity testing
negative regeneration rates occurs at a lower bound threshold, of model]
overshoot and collapse “fSR(SM)”=
occurs.As for GF3 and 5, non‐linearity has no significant impact “GF1 SM‐inflow”(Soil moisture/Initial soil moisture)
on essential behaviour (assuming zero point is maintained) Units: Dmnl
. “GF1 SM‐inflow”([(0,0)‐(1,1)],(0,1),(0.5,1),(1,0))
Units: Dmnl
SM outflow = max(Plant mass,0)*”P cons (Csm)”*”fSC(SM)”
Units: SM perc/Year

(continued on next page)

13
W. Grace Total Environment Research Themes 3–4 (2022) 100011

Soil moisture equations (continued) Herbivore equations (continued)

Soil moisture = INTEG (SM inflow‐SM outflow, Initial soil Herbivores = INTEG (H inflow + Initial H A H outflow, 0)
moisture) Units: Herbivores
Units: SM perc Multiplier
“P cons (Csm)”= 10 Units: Herbivores/Year
Units: SM perc/(Year*PMtonnes) “H‐regen‐fr (Rh)”= 0.075
“fSC(SM)”= “GF2 SM‐outflow”(Soil moisture/Initial soil Units: 1/Year
moisture) H Multiplier = 1
Units: Dmnl Units: 1
“GF2 SM‐outflow”([(0,0)‐(1,1)],(0,0),(1,1)) [Note this variable was used to vary Rh for sensitivity testing
Units: Dmnl of model]
“fHR(PM)”= “GF5 H‐inflow ‐ smooth”(zidz(Plant mass, Initial
plant mass))
Units: Dmnl
Plant mass equations “GF5 H‐inflow ‐ O&C”(
[(0,‐2)‐(2000,1)],(0,‐2),(100,‐0.85),(300,0),(630,0.55),(870,0.
8),(1600,1))
Plant mass = INTEG (PM inflow‐PM outflow, Initial plant mass) Units: Dmnl
Units: PMtonnes [Note this version of the function produces the herbivore
Initial plant mass = 25 overshoot and collapse mode of the model]
Units: PMtonnes “GF5 H‐inflow ‐ smooth”([(0,‐2)‐(3200,1)],(0,‐2),(500,0),(750,1
PM inflow = Plant mass*“PM‐regen‐fr (Rpm)”*“fPR(SM)”*PM ),(1600,1))
Multiplier Units: Dmnl
Units: PMtonnes/Year [Note this version of the function produces the herbivore
“PM‐regen‐fr (Rpm)”= 0.2 smooth or damped oscillatory mode of the model]
Units: 1/Year H outflow= “Pr cons (Ch)”*“fHC(H)”*Predators
PM Multiplier = 1 Units: Herbivores/Year
Units: Dmnl “Pr cons (Ch)”= 0.25
[Note this variable was used to vary Rpm for sensitivity testing Units: Herbivores/Predators/Year
of model] “fHC(H)”= GF6 H outflow (zidz(Herbivores,“H‐injection”))
“fPR(SM)”=”GF3 PM‐inflow”(Soil moisture/Initial soil Units: Dmnl
moisture) GF6 H outflow([(0,0)‐(1,1)],(0,0),(1,1),(100,1))
Units: Dmnl Units: Dmnl
“GF3 PM‐inflow”([(0,‐1)‐(1,1)],(0,‐1),(0.5,1),(1,1))
Units: Dmnl
PM outflow = Herbivores*“H cons (Cpm)”*“fPC(PM)”
Units: PMtonnes/Year Predator equations
“H cons (Cpm)”= 4.5
Units: PMtonnes/Herbivores/Year
“fPC(PM)”= “GF4 PM‐outflow”(zidz(Plant mass, Initial plant
Predators = INTEG (Initial Pr + Pr inflow ‐ Pr outflow, 0)
mass))
Units: Predators
Units: Dmnl
Initial Pr = 0 + STEP(“Pr‐injection”/“1 year”,200)‐STEP(“Pr‐
“GF4 PM‐outflow”([(0,0)‐(1,1)],(0,0),(1,1),(1000,1))
injection”/“1 year”,201)
Units: Dmnl
Units: Predators/Year
[Note this equation was used to introduce Predators to the
model at t200]
“1 year”= 1
Herbivore equations
Units: Year
“Pr‐injection”= 10
Units: Predators
Herbivores = INTEG (H inflow + Initial H A H outflow, 0) Pr inflow = Predators*“Pr‐regen‐fr (Rpr)”*“fPrR(H)”*Pr
Units: Herbivores multiplier
Initial H = 0 + STEP(“H‐injection”/“1 year”,50)‐STEP(“H‐inje Units: Predators/Year
ction”/“1 year”,51) + STEP(“H‐spike” /“1 year”,300)‐STEP(“ “Pr‐regen‐fr (Rpr)”= 0.03
H‐spike” /“1 year”,301) Units: 1/Year
Units: Herbivores/Year Pr multiplier = 1{6}
[Note this equation was used to introduce Herbivores to the Units: Dmnl
model and t50 and then to introduce a spike for resilience [Note this variable was used to vary Rpr for sensitivity testing
testing at t300] of model]
“H‐injection”= 10 “fPrR(H)”= “GF7 Pr‐inflow‐smooth”(zidz(Herbivores,“H‐injec
Units: Herbivores tion”))
“H‐spike”= 10 Units: Dmnl
Units: Herbivores “GF7 Pr‐inflow O&C”([(0,‐2)‐(100,1)],(0,‐2),(15,0),(40,1),(100,
H inflow = max(Herbivores,0)*“H‐regen‐fr (Rh)”*“fHR(PM)”*H

14
W. Grace
Predator equations (continued)
Predators = INTEG (Initial Pr + Pr inflow ‐ Pr outflow, 0)
Units: Predators
1))
Units: Dmnl
[Note this version of the function produces the predator
overshoot and collapse mode of the model]
“GF7 Pr‐inflow‐smooth”([(0,‐2)‐(100,1)],(0,‐2),(25,0),(40,1),(10
0,1))
Units: Dmnl
[Note this version of the function produces the predator
smooth or damped oscillatory mode of the model]
Pr outflow = 0
Units: Predators/Year
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