ACS Submission Marine Ecosystem

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Emergent features in a general food web

simulation: Lotka-Volterra, Gause's law, and
the paradox of enrichment
Article in Advances in Complex Systems · December 2013

DOI: 10.1142/S0219525913500148
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Marine-Ecosystem
EMERGENT FEATURES IN A GENERAL FOOD WEB

SIMULATION: LOTKA-VOLTERRA, GAUSE’S LAW, AND THE
PARADOX OF ENRICHMENT
TED CARMICHAEL∗
Department of Software and Information Systems,
University of North Carolina at Charlotte,
9201 University City Blvd.
Charlotte, NC 28213, USA
and
Dickson Advanced Analytics Group
Carolinas HealthCare System
720 E. Morehead St.
tedsaid@gmail.com
MIRSAD HADZIKADIC
Department of Software and Information Systems,
University of North Carolina at Charlotte,
9201 University City Blvd.
mirsad@uncc.edu
Received (received date)

Revised (revised date)
Computer simulations of complex food-webs are important tools for deepening our un-
derstanding of these systems. Yet most computer models assume, rather than generate,
key system-level patterns, or use mathematical modeling approaches that make it dif-
ficult to account fully for non-linear dynamics. In this article we present a computer
simulation model that addresses these concerns by focusing on assumptions of agent
attributes rather than agent outcomes. Our model utilizes the techniques of Complex
Adaptive Systems and Agent-Based Modeling so that system level patterns of a marine
ecosystem emerge from the interactions of thousands of individual computer agents. This
methodology is validated by using this general simulation model to replicate fundamen-
tal properties of a marine ecosystem, including: (1) the predator-prey oscillations found
in Lotka-Volterra; (2) the stepped pattern of biomass accrual from resource enrichment;
(3) the Paradox of Enrichment; and (4) Gause’s Law.
Keywords: Complex Adaptive Systems; Agent Based Model; Individual Based Model;
Lotka-Volterra; simulation; food web; Gause’s law; paradox of enrichment; resilience;
population dynamics.
∗ Corresponding author.
1
Marine-Ecosystem
2 Carmichael and Hadzikadic
1. Introduction
It is becoming increasingly evident that certain stocks of fish are becoming severely
depleted, are in danger of collapse, or have already collapsed in many parts of the
world [2, 13, 29]. In order to promote more sustainable practices, it is incumbent
on researchers to strive for a clear, holistic understanding of the complex dynamics
found in marine ecosystems. In addressing this need, we take the approach that a
computer model should be as simple as possible, yet still able to reproduce known
complex and non-linear effects. Such a model can productively serve to: (1) ad-
vance our understanding of the properties found in these systems generally; and (2)
provide a baseline for research into additional complicating factors.
The literature on marine and terrestrial ecosystems is long and varied, encom-
passing both theoretical models (e.g.: [14, 25]) and empirical surveys (e.g.: [13, 23]).
Some significant differences between model results and real-world surveys have per-
sisted for years, and it has been difficult identifying fundamental principles relative
to the many complicating factors that can be found in existent ecosystems. For
example, in the early 1980s Oksanen et al. examined multiple trophic levels in a
predator-prey system using mathematical models, in order to determine whether
species population (bio-mass) is fundamentally controlled by resources - as was the
conventional wisdom at the time - or dominated by predation [39]. In describing
this work, Power states that these models produce “a stepped pattern of biomass
accrual” [42]; Brett and Goldman further characterize the Oksanen et al. results,
saying that “In food webs with an odd number of trophic levels, increases in primary
production should lead to increased biomass for odd-numbered trophic levels and
no change in biomass for even-numbered trophic levels. Conversely, in food webs
with an even number of trophic levels, increases in primary production should lead
to increased biomass for even-numbered trophic levels and no change in biomass for
odd-numbered trophic levels” [7].
More recently, researchers have begun to ask, not which process (bottom-up
resources vs. top-down predation) dominates overall, but rather how these two
forces interact under different spatial and temporal scenarios [43]. Many scientists
think that this new approach will help reveal “how resilient food webs are to the
combined effect of resource-mediated and predator-mediated forces” [11]. This sug-
gestion leads naturally to the view that ecosystems can be productively studied as
Complex Adaptive Systems; that is, systems that are characterized by their emer-
gent properties, self-organization, and non-linear dynamics [1, 8, 24, 28, 49]. The
field of Complex Adaptive Systems (CAS) recognizes that these systems are gener-
ally robust and flexible, consisting of multiple negative feedbacks that produce one
or more “basins of attraction:” i.e., the emergence of resilient, system-level patterns.
Indeed, it is the identification of the key feedbacks and their contributions to system
resilience that is the ultimate goal of ecological research.
Advances in computing technology have allowed for more robust in silico sim-
ulations that can model these complex ecosystem dynamics in ways previously in-
Marine-Ecosystem
Emergent Features in a Food Web Simulation 3
tractable. In order to address certain limitations of purely mathematical models

[14, 41], which generally simplify individual variation and spatial representation,
and also tend to represent global properties in a top-down manner, we have created
a general CAS model of a marine ecosystem. This model reproduces classic phenom-
ena of population dynamics, but in a stochastic, agent-based computer simulation,
so that the complex dynamics of predator and prey interactions can be more fully
explored.
Central to a more flexible, Agent-Based Modeling (ABM) approach (sometimes
called Individual-Based Modeling [12], or Agent-based Complex Systems [26]) is
that the outcomes of the simulation are generated in a bottom-up design process,
rather than via top-down constraints [16]. It is important to note that the phrase
“bottom-up design” has a somewhat different meaning in the CAS literature than
the similarly-worded “bottom-up” forces referred to in marine and terrestrial ecosys-
tem literature. A CAS-based ABM implies that the system-level patterns, such as
population growth, aggregate predation rates, etc., are generated from the bottom
up, rather than assumed as a top-down constraint on the modeled system [20]. That
is, the system-level patterns are emergent properties that arise from the interactions
of the autonomous agents that comprise the simulated system. Thus, the method
of computer simulation relies on assumptions of agent attributes rather than agent
outcomes. (The similar phrasing in the ecosystem literature is unfortunate; one
might come across, e.g., a discussion of the controlling forces in constraining a par-
ticular population: i.e., “bottom-up” forces of resource availability vs. “top-down”
predation. This is an entirely different context than “bottom-up” emergence in an
ABM.)
Here we present a general ABM of predator-prey interactions with a focus on key
phenomena in population dynamics. This methodology is validated in a number of
ways. The first is by replicating the cyclical relationship between the predator and
prey populations as predicted by the classic Lotka-Volterra (LV) equations. Sec-
ond, this model reproduces the stepped pattern of biomass increase that Oksanen
et al. predicted through the use of mathematical modeling. Third, the model also
reproduces the “Paradox of Enrichment” effect: that is, under multiple simulation
scenarios, increasing primary production leads to increased volatility [46–48]. Fi-
nally, our simulation model also exhibits the Competitive Exclusion Principle, also
known as Gause’s Law, which states that two or more species that compete for the
same resources in a constant environment cannot co-exist.
2. Materials and Methods

2.1. Complex Adaptive Systems
A Complex Adaptive System (CAS) is a collection of autonomous, heterogeneous
agents, whose behavior is defined with a limited number of rules. These rules gov-
ern the type and number of interactions among agents. The power of the system
mainly comes from agents’ interactions, not the agents themselves. Each individual
Marine-Ecosystem
interaction generally has only a small or limited direct effect on the outcome of the
system. However, the aggregate product of the thousands of these interactions and
the accumulated feedbacks among the agents can have a large effect.
CAS is a method developed in physics, mathematics, and computational sciences
[6, 24, 28, 31, 35, 38] to deal with the issue of complexity and complex systems,
and has been redefined by a growing number of applications in many domains.
Complex, dynamical systems are comprised of parts that interact with each other.
They are complex because it is difficult, if not impossible, to predict their behavior
by merely understanding the function of each part. The diversity of these parts
and the richness of their interactions endow a complex system with its capacity
to innovate, adapt, and sustain itself. At the same time, these global, emergent
properties cannot be studied or readily understood by only inspecting the parts in
isolation.
Agent-Based Modeling (ABM) is a method for implementing a CAS in order to
provide a computational environment for exploring complex systems in a controlled
setting. Since the mid-20th century there has been a steady effort to apply CAS
to areas as diverse as economics, business, political science, government, military,
archeology, biology, and ecology [3–5, 9, 10, 17, 21, 22, 27, 30, 33, 34, 36, 37, 40,
44, 45, 50]. Designing CAS applications is challenging because researchers often do
not know what key variables need to be captured to successfully model the system.
There has been some evidence that the CAS method itself can be used to identify
key system variables [19].
A generative ABM approach has a number of advantages. Our emphasis on
bottom-up design principles ensures a more robust simulation, in that such a model
can allow simulations to have a wide degree of flexibility through their ability to
generate emergent behaviors. An ABM is also inherently more representative of
true biological systems, because the simulated agents are analogues to their real-
world counterparts. Further, this property allows for easy exploration of alternative
theories and/or assumptions, simply by changing a few of the agent attributes or
rules of behavior. Such flexibility and transparency is important for exploring the
parameters of known phenomena, as well as for calibrating the ABM to specific
empirical data.
The generative nature of the system-level phenomena via bottom-up processes
ensures that the complex dynamics at work in the simulation are much more accessi-
ble than with traditional, top-down simulation design. For example, a “black-box”-
type simulation will often use complicated mathematical formulae to constrain the
system. There may even be multiple, competing functions of this type that define
the system properties. The way in which these functions interact can compound
the difficulty of deciphering how they produce the simulation outputs. Conversely,
a bottom-up ABM design relies only on agent-level assumptions; the system-level
properties are allowed to emerge primarily, or solely, from the interactions of these
individual agents.
Although it is not always clear how some emergent features are connected to the
Marine-Ecosystem
agent behaviors, this is also true for many real-world situations. Consider, for ex-
ample, China’s “one-child” policy. This is an agent-level policy, aimed at producing
a particular system-level result (lower population growth). The policy goal (emer-
gent feature) must be translated into agent-level rules of behavior. Even though the
exact results can be hard to predict, the policy itself is obvious and transparent.
A bottom-up ABM is the same, with agent-level assumptions rather than system-
level assumptions. Further, in such an ABM one can easily experiment with different
agent-level rules and attributes, to explore potential outcomes. Thus the assump-
tions in this design method are inherently transparent, giving the model advantages
in terms of both flexibility and in understanding. A black-box-type simulation, such
as when the component functionalities are obscured, or particular controlling algo-
rithms are represented by complicated mathematical formulae, does not have these
advantages.
We created our marine system simulation using the NetLogo programming en-
vironment. NetLogo is a general ABM simulation tool ideally suited for rapid-
prototyping of complex systems [51].
2.2. Lotka-Volterra
First proposed in 1925-1926, the Lotka-Volterra (LV) equations are a pair of first-
order, non-linear differential equations that govern the relationship between two
types of interacting species. The equations have periodic solutions, such that an
increase in the prey population generates a temporary increase in the predator pop-
ulation, which increases predation levels. Increased predation reverses the growth
of the prey population, which in turn reduces the predator population. Once the
prey reverses again to a growth phase, the cycle is complete.
Let x equal the prey population and dx equal the rate at which the prey pop-
ulation increases. In the expression of LV, dx = Axdt, where A is the growth rate
of the prey population and dt is the change in time t. Thus, in the absence of some
predator the prey population x will increase exponentially.
We define y as the predator population, and dx = -Bxydt as the rate of prey
consumption by the predators. Thus the overall growth rate in the prey population
is given by combining these two equations:
dx
= Ax − Bxy (1)
dt
For the predator population, overall growth is dependent on C, the rate of preda-
tor removal from the system (either by death or migration), and D, the growth rate
for predators. Predators therefore also have two equations to express the change in
population over time: 1) dy = -Cydt and 2) dy = Dydt, which combine to form the
following equation:
dy
= −Cy + Dxy (2)
dt
Marine-Ecosystem
The solution to these two equations is periodic, with the predator population
curve always following the prey curve. Figure 1 gives a graphical example of typical
LV solution (left image), and displays the same results - but in a more stochastic
way - as seen in our simulation model (right image).
Fig. 1. Example of a periodic solution for the LV equations (left) and the stochastic behavior of
predator and prey populations in our general ecosystem simulation (right).
2.3. Modeling Methodology

For instantiating a general marine ecosystem model, a number of controlling as-
sumptions of the LV equations have been adopted, while others have been changed
to allow for greater flexibility in the model. The key assumptions of LV are: (1)
unlimited food availability to the prey population; (2) the predator population
depends entirely on the prey for food; (3) the natural growth rates for both pop-
ulations are proportional to their sizes; and (4) the environment does not change
to the benefit of either population. Other fundamental assumptions of LV include
homogeneity of the agents and the system, and that the species are not adapting
to their environment.
Our investigation into a deeper understanding of the predator-prey dynamics
began by changing assumption (1) and (3) above, which are connected. Assumption
(1) states that the food is unlimited, and thus both the predators and the prey can
always consume, and reproduce, at a rate that is relative only to their population
sizes (assumption (3) above). These rates are expressed in the LV equations with the
coefficients A and D: A is the growth rate for the prey population, and, therefore,
Ax is this rate times the prey population size, x; D is the growth rate for the
predators, and Dxy is the product of this rate times the prey population size, x, and
the predator population size, y.
These assumptions inherent in LV demonstrate how an ABM can be useful.
Rather than assume constant consumption/reproduction rates (A and D), we can
instead force our simulated agents to find food. As with LV, the rate is still propor-
Marine-Ecosystem
tional to population size. For the prey population, the maximum rate is to eat once
per simulation time-step; however, if other prey eat all the local food first, then
this rate is less than maximum for that particular individual. Thus the growth rate
for the prey population is constrained by the growth rate of food and by compe-
tition, rather than artificially constrained by some variable, A. The predators also
eat only once per time-step. Similar to the prey, if they cannot find food then they
dont eat. Thus the predator population growth rate is directly constrained by both
the growth rate of the prey and by competition; and indirectly constrained by the
growth rate of food for the prey. In this way, the ABM expression of both A and
D are still proportional to population size, but in a more realistic and stochastic
manner, allowing for the possibility of a variable growth rate.
The simulation environment is a torus grid with 151*151 grid cells, for a total of
22,801 cells (or “patches” in NetLogo terminology). The main simulation window is
shown in Figure 2. There are four populations in the baseline model: food (generated
by the simulation stochastically); fish (the prey population); fish-eggs (generated
by the fish as a positive function of the amount of food consumed); and predators
(which reproduce as a positive function of the number of fish consumed). We ran
additional experiments with a fourth trophic level, labeled top-predators, which
reproduce as a positive function of predators consumed; these results further confirm
Oksanens stepped pattern of biomass accrual and are discussed in Appendix A.
This model is not intended to include all the properties of an existent ecosystem,
but rather to expose the most fundamental properties of the predator-prey-resource
relationship as a general model. As such, the environment is largely homogeneous:
that is, there are no variations in sea temperature, depth, or ocean currents. Further-
more, each trophic level is represented by a single species. Important refinements -
such as species growth over time, variable predation strategies, environmental het-
erogeneities and dynamics, more complex food web networks, and different func-
tional responses - can be selectively added to future models in an iterative process
to ensure that one understands the basic dynamics at each level before proceeding
to the next level of complexity.
As with the environment, both the fish and the predator populations are homo-
geneous, different only in their current state variables: individual age, x-y coordi-
nates, and a counter to record how much food has been consumed. They are also
homogenous in terms of their simulated functions, and - unlike in the real world -
there are no simulated changes based on age or growth.
Other simplifying assumptions include: all species are the same size, produce
the same amount of resources when consumed, and share the same set of simple
strategic rules. For the fish these rules are: if there is one or more predator on
the current patch, pick one and move in the opposite direction; if no predators
are present and there is food on the current patch, eat one unit; otherwise, move
randomly. For the predators the rules are: if there is one or more fish on the current
patch, eat one. After eating (or not, if no fish are present), move randomly.
The fish, predators, and top-predators all have a limited lifetime. Also, the
Marine-Ecosystem
Fig. 2. Image of the marine ecosystem simulation environment, developed using the NetLogo envi-
ronment. Controls for each of the four populations (fish, fish-eggs, predators, and top-predators)
allow the simulation operator to change various parameters, such as speed, resources required to
spawn new agents, lifetime, etc., including environmental (“patch”) parameters. The main window
(right side, top) gives a graphical/spatial display of the currently existant agents. Two plots are
shown here, which output the population levels (left) and average age (right) of the fish and the
predators. Additional plots, controls, and monitors (not shown) display various output measures
such as current consumption rates, reproduction rates, exact population sizes, and others.
patches grow food (resources for the fish) stochastically, based on an operator-
controlled setting that defines the percentage chance of growth for each individual
patch. Food is simulated as units per patch, from zero (no food) to a maximum,
such that if food is present a fish can eat one unit per turn. Specifically, a 0.20
food growth rate translates into a 20% chance for each patch to add one unit of
food, during each simulation time step. Aggregated across the 22,801 patches in the
simulation, this rate becomes a linear, but still stochastic, rate of growth for food.
In all experiments reported here the maximum number of food units per patch is
Marine-Ecosystem
set to ten.
The population of fish eggs is included to provide another step towards a more
realistic simulation; however, all major results listed here - LV oscillations, Oksanen
et al.’s stepped pattern of biomass accrual, the paradox of enrichment, and Gause’s
Law - are reproducible whether the simulated fish produce eggs or, as with the
predators and top-predators, spawn asexually.
The majority of experiments are run with a baseline model from which any
experimental deviations are made. As the classic LV oscillations, as shown in Fig-
ure 1, are inherently volatile, we adjusted certain parameter settings in order to
de-emphasize this behavior and produce a more stable system from which to exper-
iment. This allows us to compare and contrast results under stable, oscillating-but-
stable, and unstable settings, as well as expose which settings produce each.
3. Results
3.1. Stepped Pattern of Biomass Accrual
When we run the simulation with a baseline test-case (food-fish-predator; food
production set to 0.20 chance of positive growth) it settles to an equilibrium rela-
tionship between the fish and predator populations. (See Appendix A for results
from the four-trophic-level simulation.) The exact numbers are quite stable, but
are themselves immaterial, as they would differ under different species settings and
attributes. What is of interest are the changes in these numbers, however, and what
these changes demonstrate in terms of the inherent dynamics of increasing the food
supply.
In terms of age, the equilibrium age for predators is about 50% higher than that
of fish. In Figure 3, we see how variations in the food supply - the lowest trophic
level - affect the relationship between the mid- and high-trophic level populations.
The left graph in Figure 3 shows the change in population for the fish, eggs, and
predators, as the food supply is increased from 0.20 to 0.30, and again to 0.40. The
graph on the right displays the resultant change in average age for these populations.
Initially there is a fish population “spike” in response to an increased food supply.
The predator population then increases in response to this extra prey. Remarkably
however, the gains in fish population are only temporary, and are quickly offset
by the increased predation rates. Thus, the fish population returns to the same
equilibrium level as that found with a lower supply of food: only the predator
population remains elevated. This result indicates that all the gains stemming from
the increased food supply accrue to the high-trophic-level predators.
When we examine the results of the average age for each population, we see
that the fish - though reproducing at a faster rate - do not live as long as they did
before. Even as they reproduce faster, they are also consumed faster, so that their
average age is much lower. This is the necessary consequence for preserving the
equilibrium prey population noted above. The predators, faced with an increased
food supply, are initially much more efficient in catching the fish; thus, their pop-
Marine-Ecosystem
Fig. 3. Plotted outputs after 3000 simulation time-steps, showing population sizes at 20%, 30% and
40% food growth levels (left image) and the resultant change in average age for each population
(right image). When the food level is increased after 1000 time-steps to 0.30 - and again, at 2000
time-steps, to 0.40 - there is an initial dramatic rise in the fish population. However, once the
predator population increases in response, the fish population returns to approximately the same
size as before.
ulation increases. Unlike the fish however, their average age - after stabilizing at a
higher population - is essentially unchanged. Table 1 shows the predator and prey
consumption rates under different rates of food growth, after stabilizing at each new
level. This table illustrates that, even though the fish population does not increase,
their rate of activity (how fast they consume food) does. The predators, however,
have no persistent benefit in consumption rates: the have more fish to eat, but this
is balanced by increased competition (more predators).
Table 1. Predator and fish consumption levels.
Enrichment Level Predator Consumption Rate Fish Consumption Rate

0.20 food growth 0.049 0.244
0.30 food growth 0.049 0.364
0.40 food growth 0.048 0.485
0.50 food growth 0.049 0.600
Note: Consumption rates for fish and predators for four different levels of
resource growth (0.20, 0.30, 0.40, and 0.50) calculated over 1000 simulation
time-steps. The consumption rate is the average number of units of food
consumed per time step, per individual of that species.
Though other factors will ultimately influence these dynamics to some degree,
the overall picture is clear: the highest trophic level - the predators - is greatly
affected (in terms of population levels) by the food available at the lowest trophic
level, while the species in the mid-trophic level - the fish - is affected only in average
age and reproductive activity, not in population size.
Marine-Ecosystem
3.2. The Paradox of Enrichment

The previous results revealed what happens when the lowest trophic level - the fish
food supply - changes from 0.20, to 0.30, and again to 0.40. Another important
question is: what happens when this food supply is continually increased, even be-
yond 40%? Figure 4 shows the results as the food supply is incrementally increased,
from the initial 20% as before, with 1% added each 100 simulation time steps.
Fig. 4. Plotted outputs after 3600 simulation time-steps. Food supply is increased by 1% (0.01)
every 100 time-steps, from 0.20 food growth to 0.55. The left-side panels display the relative
population levels (top) and the average population ages (bottom). The right-side panel displays
a representative image of the spatial characteristics: i.e., the x- and y-coordinates of each agent
in the simulation, as well as the current state of the 22,801 patches. For the patches, light grey
indicates low levels of existent food while dark grey/black indicates high levels of food.
One can see that there is a marked non-linear change in volatility, here at ap-
proximately the 42% food growth rate. (Again, only the presence of a threshold level
for instability is significant, not the exact level it emerges, as this level would change
under different parameter settings.) At this point the increased food still seems to
translate into only the predators becoming more populous. The prey population,
while much more volatile than the predators, nevertheless oscillate around a fairly
stable level. As before, the predators are holding fairly stable in terms of average
age, while the fish tend to die more quickly as their food increases. However, we
also see that the average age of the fish does not decrease at a linear rate. That is,
the fish average age seems to be approaching some minimum, below which it cannot
reach. This may account for the increased volatility in their population numbers.
Spatially, the simulation environment is becoming more heterogeneous at the
point where volatility suddenly increases. Up until the critical point of 0.42 food
Marine-Ecosystem
growth, the environment indicates minimal variability, and the four populations are
well-mixed throughout. At the same point where population-size volatility begins,
the spatial relationships between these populations also change: a purely endoge-
nous effect in the simulation environment. In some places, food is allowed to grow
unconsumed, while in others it is completely wiped out, in a pattern that appears
and disappears across the simulation environment over time (Figure 4, right image).
The volatility of the populations becomes even more pronounced as food levels
are continually increased - an increasingly non-linear response to nutrient enrich-
ment. Also, the fish population, which had previously oscillated around a stable
amount, now seems to be increasing somewhat. The dramatic swings between the
high and low population levels are much more extreme. As previously noted, these
seems to occur as the fish population average age reaches a minimum that cannot
be crossed.
The increased volatility is problematic for both the fish and predator species.
Due to the stochastic nature of the ABM, and the spatial clustering of the pop-
ulations, there is an increased chance that one of the populations will completely
die out. In this sample run, this occurred at approximately 4200 time steps, at a
food growth rate of 0.61 (not shown). Repeated simulation runs confirmed that this
approximate level is indeed unsustainable under these settings.
As with the total population numbers, the volatility of consumption rates for
both predator and prey increase also. Comparing Figure 5 with Figure 4, one can
see the relative volatility of consumption rates with that found in average age and
population levels.
The outputs of the simulation allow us to directly measure key parameters of
both fish and predator growth, predator consumption of fish, and the rates that
both species die due to limited lifetime. Further, one can see how each of these
parameters is affected under different scenarios of food growth rates at the lowest
trophic level. Table 2 shows these results for 0.20, 0.30, and 0.40 food growth rates.
Table 2. Consumption rates, species additions, and species re-

movals.
Nutrient growth rates: 0.20 0.30 0.40
Fish Consumption 0.244 0.364 0.485
Predator Consumption 0.049 0.049 0.048
New fish per time step 73.6 144.1 223.9
New predators per time step 2.2 4.9 7.6
Fish death rate (due to old age) 9.3 1.6 0.2
Predator death rate (due to old age) 2.2 4.8 7.6
Note: Fish and predator consumption rates (taken from Table

1), new fish and predator rates per time step, and rate of agent
death due to limited lifetime per time step, for three different
levels of resource growth: 0.20, 0.30, and 0.40. These numbers
are averaged across 1000 simulation time steps.
Marine-Ecosystem
Fig. 5. Consumption rates over time for both the predator population (top left panel) and prey
population (bottom left panel) as food growth is slowly increased from 0.20 to 0.55 (3600 time
steps; note that the y-axis ranges from 0.0 to 0.3 for the predator consumption rate, and from 0.0
to 1.0 for the prey). When compared with the outputs found in Figure 4 (reproduced here, right
two panels) one can see that volatility in consumption rates occurs at the same time volatility
emerges in the population levels.
Note that the LV equations do not account for prey dying from old age, or oth-
erwise being removed from the system; they can only be consumed by predators.
As their average age decreases, however, this difference from LV becomes nearly
non-existent, because the number of fish that die of old age approaches zero. The
predators death rate, meanwhile, approximately equals the new predators intro-
duced into the system. (Due to the stochastic nature of the simulation, these two
rates should not always be exactly equal.) This is what one would expect, since the
predator population is quite stable.
3.3. The Competitive Exclusion Principle

The competitive exclusion principle, also known as Gause’s Law, is a well-known
mathematical result in ecosystem dynamics. Simply stated, two or more species that
compete equally for the same resources in a constant environment cannot co-exist.
This law is problematic, however, as the predicted result seems to seldom occur
in nature. This state of affairs is thus known as the Paradox of the Plankton, a
long-standing conundrum in ecological research.
Nevertheless, we tested our model, under multiple simulation runs, and under a
variety of nutrient enrichment scenarios in the baseline model, and have confirmed
Gause’s Law for the prey population in the three-tropic-level simulation. For these
Marine-Ecosystem
runs the fish population is divided into two distinct species. Neither species possesses
any functional difference or advantage, and they are distinguished in the simulation
only by color. We ran thirty simulations for each of 0.20, 0.30, 0.40, and 0.50 nutrient
growth. In these simulation runs, one of the two species of fish would generally die
out within ~1000 simulation times steps. (For context, the average age of the fish is
~80 times steps at 0.20 food growth, and less - consistent with the results in sections
3.1 and 3.2 - for higher levels of nutrients.) In general, the mean number of time
steps required for this to occur was slightly higher for increased level of nutrients,
reflecting the elevated level of fish consumption and reproduction activity. (See
Appendix B for more details of these results.)
4. Discussion
For the LV equations, the first major difference is that LV assume a “natural” level
of population growth: a rate that is proportional to the size of the population. In
the equations outlined (Eq. (1) and (2)), recall that term A is the growth rate for
the fish, and D is the growth rate for the predators. Both of these terms are assumed
to be constant in LV.
In our model, however, these terms are stochastic, dependent upon consumption
rates Unlike the assumptions underlying the LV equations, neither the fish nor the
predators are guaranteed to find food each time step. For the prey in the three-
trophic-level simulation we find that the fish consumption rate, and subsequent
reproduction rate, increases with resources. As food grows more on the patches (with
the increased rate of growth), the fish have an easier time, and thus reproduce more.
This is tempered by the fact that, as the fish population increases, the predators
subsequently have an easier time finding food.
The consumption rate of the predators eating the prey, coefficient B in the
LV equations, is also not constant. However, their consumption rate only increases
when the system is in transition between one stable state and another. Once the
predator population reaches a new level, the aggregate consumption rate returns to
the equilibrium rate as seen in Table 1.
The growth of the fish is also no longer held constant. In this simulation, the
variable nature of A is not only due to how much food is present locally, but also
due to the presence of predators locally. If a fish sees a predator, it moves away
without eating, even if food is present. Further, local competition among the fish
can also affect A: as the number of fish increases, finding food - even in the absence
of predators - becomes more difficult. The spatial distribution, therefore, becomes
quite important.
These departures from the LV equations can help explain the volatility that ex-
ists at higher levels of resource growth - the Paradox of Enrichment in the ecosystem
literature. At higher predator densities, the fish often die too early to reproduce,
even when there is plenty of food. They will begin to congregate on “safe” patches,
while food will grow unmolested on “unsafe” patches. This behavior, though en-
Marine-Ecosystem
tirely the result of random movement and spatial distributions, generates the en-
dogenously created heterogeneity that is seen in Figure 4.
The final departure from LV is that both the fish and the predators in this
simulation have a lifetime. The LV equations account for a lifetime for the predators
only, via the C coefficient. (Recall that the C coefficient is the rate in which predators
are removed from the system, either by death or migration.) In our model, we
simulate the rate by tracking the age of each agent, coupled with a limited lifetime.
As noted previously, the simulation’s emergent value for C remains largely constant,
even under different population levels of predators.
For the prey, LV doesn’t remove individuals from the system based on age or
migration, but our simulation does. This allows for additional insight into the causes
of increased volatility at certain levels of resource growth. We have seen that the
average age of the fish decreases - and consumption/reproduction activity increases
- when food levels go up. But there is a minimum average age below which the fish
simply cannot get any younger. Intuitively, one would think that an increase in fish
too young to reproduce would be bad for the fish. But in fact, it is bad for the
predators, because they still depend on the fish for their food. It is at this point
that the predation begins to lose its effectiveness in regulating the fish population,
and the dangers inherent in the Paradox of Enrichment become manifest.
In terms of Oksanen et al.’s stepped pattern of biomass accrual, the model vali-
dates the finding that, in the three-trophic-level simulation, all increases in the re-
source availability accrue to the predator species, and none (excepting slight stochas-
tic variations) to the prey. Further, in the four-trophic-level simulation (Appendix
A), this accrual of resources is evenly divided between the prey (level two) and the
top-predators (level four), with none going to the level-three predators. Although
this agrees with Oksanen et al., the ocean survey literature is divided on this point,
with evidence from nature sometimes supporting this result and sometimes not.
Nevertheless, our results confirming Oksanen et al. represent a generative,
baseline-case dynamic that can be used to judge complicating factors found in
existent ecosystems. Importantly, there are two clear quantitative comparisons that
one can make: total biomass of each species (in the simulation, this is a simple
function of population size); and the increase in activity/decrease in average age
of the prey species. The first observation, that the predator population increases
when resources for the fish are increased, may at first seem to contradict the fact
that the predators’ consumption rate does not change. More predators would imply
that there is more competition among them, and thus seem to warrant a lower con-
sumption rate. However, the increased resources for the fish, as we have seen, help
produce more fish that are consumed more quickly. Therefore, as the number of
predators increases, so does the availability of prey. Remarkably, these two changes
seem to balance out perfectly and, therefore, competition among the predators re-
mains stable under multiple resource levels.
In terms of Gause’s Law, our results confirm the competitive exclusion princi-
ple for the prey population. Recall that we adjusted this model to include two fish
Marine-Ecosystem
species, distinct but functionally identical. Under the four scenarios of varied nutri-
ent levels across 120 simulation runs, the greatest number of time steps needed for
one of the two fish species to die out was ~2300, and the mean run time was ~1240
steps. In this scenario - 0.50 food growth - the average age for a fish individual is
generally 30-34 time steps. If one assumes a generational length equal to mean age,
then this length of time represents less than 40 generations before one of the two
species becomes extinct.
As with any model, our ABM reduces the complexity of the real world by strip-
ping away complicating factors, such as the many heterogeneities found in nature.
Thus greatly simplified, the ABM nevertheless reproduces the idealized mathemat-
ical result found in Gause’s Law. And, in fact, the changing population levels of the
two fish species, as their levels wax and wane, is reminiscent of a “random walk.”
Eventually, one of the two populations will cross the zero line, but it is impossible
to predict which one, or precisely how long it will take.
As previously noted, Gause’s Law is a purely mathematical result, one that
is almost never found in nature. That is, there are a multitude of examples where
multiple species do seem to compete for the same resources, in a stable environment,
and yet manage to persist. This antithesis to Gause’s Law is known as the Paradox
of the Plankton, and is called a “paradox” due to being at odds with the clear
mathematical consequence of fair competition.
Many hypotheses have been put forth to explain this paradox, including external
factors such as gradients of resource availability; dynamic fluctuations in water tem-
perature, turbidity, or depth; or differential predation. Multiple species that exist
at approximately the same trophic level may have subtle differences in efficiencies
or resource requirements. Spatial heterogeneity may be created endogenously, as
we saw in Figure 4, or due to other endogenous or exogenous effects not simulated
here.
Another possibility is that the answer is more subtle, and that diversity in some
systems is long-term unstable, but “long-term” is on a scale that is so large as to
be immaterial. John Maynard Keynes’s famous statement, “In the long run we are
all dead,” captures this idea in the context of economics, but could just as easily
apply to complex ecosystems [32].
Nevertheless, the debate on the paradox of the plankton and associated issues
regarding which characteristics of food webs promote stability is a long running one
(see, e.g.: [1, 49]; for a review of the evolving debate on food web networks, see:
[18]). Therefore, in building a simulation model that can be used to explore these
issues, it is crucial to first be able first to reproduce the mathematical principle of
Gause’s law with a generative simulation. Only then can subsequent refinements,
added one at a time, be used to explore how diversity may emerge and persist, and
under what conditions it does so.
Marine-Ecosystem
4.1. Conclusions and Future Work

The simulation model presented here represents a realization of the classic LV equa-
tions in the form of an ABM. The model design is purposefully built to be a general
representation, yet one that is capable of more realism and flexibility than intended
by the idealized LV equations. For example, the agents - both predator and prey -
have a limited lifetime. They also have consumption rates (along with the related
rates of reproduction) that are stochastic: dependent on the spatial distribution
of agents rather than fixed values in a formula. While this model has a degree of
greater realism, there are many factors in a true marine ecosystem that have not
yet been factored in. This was done purposefully, so that the complex dynamics of
this system could be fully explored before additional complications are added to the
model. It also highlights the utility of the ABM approach for simulating complex
dynamics, in that the additional flexibility afforded here is inherent in this method.
In general, the additional factors that we plan to add to future iterations of this
model are ones that make either the environment or the agent populations more
heterogeneous. For the environment, it may be useful to allow for variation of such
factors as ocean depth and temperature, and ocean currents. For the agents, one can
add greater heterogeneity by allowing each species to grow, perhaps moving from one
trophic level to another. Also, real-world ecosystems usually contain multiple species
at each level, rather than just one, as well as a complex food web of dependencies
and interactions, including more and varied predation links among species. Further,
each member of these species would have some variation in their attributes, unlike
these simulated agents, such as how fast they can move, how much food they require,
and a varying ability to escape predators.
Finally, we note that the direct relationship between consumption of food and
reproduction is not as clear-cut as represented in this model. While it seems clear
that the lack of food in a real ecosystem would be a limiting factor on population
growth, it is not as clear to what degree an excess of food has on a linear assumption
of reproduction. This is also where considerations of seasonality would presumably
make a difference as well, as many species of fish reproduce only during certain
times of the year, or in certain places.
Nevertheless, this model’s primary results provide a useful baseline reference
of fundamental dynamics. Further, the effects of resource accrual towards higher
trophic levels could have many practical implications for the management of ocean
fisheries. For example, some fishing techniques include a method whereby the fish-
ing nets are anchored on the seabed. As the net is dragged along behind the ship,
the seafloor in this area is completely disrupted, and decades worth of accumu-
lated marine structures are destroyed. Previous generations of fishermen may have
assumed this does not much affect the species being harvested, due to the target
species generally existing one or more levels away from these low-trophic-level life
forms. However, if a robust ecosystem at the lowest trophic level can accrue biomass
all the way up to the highest trophic level, then such fishing practices may in fact
Marine-Ecosystem
represent twice the disruption to their robustness: reducing the current numbers
via direct harvesting, and reducing their future numbers by undermining the fish’s
ultimate source of food.
Any simulated relationships and dynamics have to eventually be validated with
real-world empirical evidence. Yet by simulating these species in an ABM, we gain
much in terms of understanding the fundamental dynamics of these systems. Thus
this model allows a level of accessibility to core assumptions and activity - across
multiple species and multiple trophic levels - to a degree that is often impossible in
real-world surveys. Further, additional steps towards a more realistic simulation can
be added in ways that might be impossible to find in empirical studies. This property
is crucial in particular for future exploration of Gause’s Law and the Paradox of the
Plankton, as this paradox has many competing potential solutions in the literature.
In fact, it may be the case that there are multiple solutions to the paradox; i.e.,
there is likely more than one single answer that can explain how diversity is able to
persist in nature. Since this model can reproduce Gause’s Law, it can now be used
to experiment with added complications, singly or in combination, to find under
what conditions Gause’s Law is not true.
The possibilities for understanding fundamental processes and dynamics via
this model is not restricted, as is done here, to marine ecosystems. A natural ex-
tension would be to consider terrestrial population dynamics as well. Further, the
LV equations themselves were originally developed to describe autocatalytic chem-
ical reactions, and we see no reason why this ABM instantiation of these equations
would not also prove useful in chemistry, with appropriate metaphorical mapping
to that domain. We also note that LV has been applied to economic systems as far
back as 1967 [15]. We are intrigued by the possibility of Oksanen’s stepped pat-
tern of biomass accrual being applied to economic theory. There are several hurdles
to be overcome, such as drawing useful analogies that address the distinction be-
tween prey (that produce offspring to be consumed) verses companies (that produce
goods and services). Nevertheless, such a model may introduce a fruitful “trickle
up” economic theory, such that the “predators” - the largest or richest entities in an
economic model - benefit due to increased resource allocation at the lower trophic
levels (the smallest or poorest entities) even if, as in Oksanen et al., this benefit
only occurs indirectly.
Acknowledgements
The authors wish to gratefully acknowledge Paul Youngman for proofreading this
article and providing comments.
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Appendix A. Stepped Pattern of Biomass Accrual - Extended

Report of Results
The main article reports the results of the three-trophic-level simulation and its
agreement with Oksanen et al. [39]. As discussed in the main text, Oksanen et al.’s
predicted results is qualitatively different for a four-trophic-level simulation. Here
we show that our model agrees with this prediction as well.
We adjusted the model to add another species, which we label “top predator.”
The top predator species consumes its prey and reproduces in the same way as
the predator species; i.e., top predators search for and consume predators in the
same way that predators consume fish; and their reproduction is a linear function
of the number of predators consumed. Other model settings were adjusted slightly
to present these results within a stable range; i.e., outside of the increased volatility
found with elevated nutrient levels. Figure 6 shows, as with section 3.1 in the main
text, the results at 0.20, 0.30, and 0.40 nutrient levels.
Fig. 6. Illustrative results from a four-trophic-level simulation. The fish (prey) species is plotted
in light grey, the predator species in medium grey, and the added “top predator” species is in
black. Nutrient levels of 0.20, 0.30, and 0.40 for 1000 time steps each. The left graph displays the
population level (top predator population is scaled by ×10 for easier comparison) and the right
graph displays the average age for each population.
Marine-Ecosystem
As one can see, this result is significantly different from that found in the three-
trophic-level simulation. As predicted in [39], here the prey species (level 2) popu-
lation and the top predator (level 4) population both increase with higher levels of
nutrients; further, the predator (level 3) population remains the same throughout.
Similar to the results of the three-trophic-level simulation, the predator population
remains at a constant level even though they consume more fish with increased
nutrient levels. In the right-side graph we again see that the population can remain
stable due to a decrease in the predators’ average age. The predators eat more,
and reproduce more, but they are themselves eaten faster by the top predators;
therefore they have a shorter average lifetime and, hence, a stable population level.
Appendix B. Competitive Exclusion Principle - Extended Report

of Results
Results from multiple runs exploring the competitive exclusion principle. In Figure
7 thirty simulation runs for each level of enrichment (0.20, 0.30, 0.40, 0.50) are
shown. The prey population was replaced with two distinct “fish” species, each
functionally exactly the same (no advantages or disadvantages based on speed,
nutrients, strategy, etc.) These were differentiated and tracked only by their color,
so that ”red” fish would spawn new red fish, and ”green” fish would spawn new
green fish. Both species consumed the same resources and are consumed without
preference by the predator species.
Fig. 7. Graphical representation of 30 simulation runs, displaying the time (in simulation time
steps) elapsed before one of the two prey species dies out. Results are shown for nutrients levels
of 0.20 (top left), 0.30 (bottom left), 0.40 (top right), and 0.50 (bottom right).
At initialization, fish were randomly assigned to ”red” or ”green.” The sim-

ulation runs were stopped when one of the two populations reached zero. In all
Marine-Ecosystem
simulations this eventually occurred. Table S1 shows the mean number of simula-
tion steps required for one of the two populations to reach zero. (As expected, this
process takes slightly longer for higher levels of enrichment.) Figure 7 graphically
shows the number of steps for each run. Initial conditions are unable to predict
which of the two species will eventually die out.
We also completed a number of experiments on the competitive exclusion prin-
ciple in regards to the predator population, formulated in the same way as with the
prey (i.e., two, and then three species of predators, but with no functional differ-
ence between species.) Generally, we find that the trend is similar as with the prey:
that one species only will remain, and the relative populations between the two (or
three) displays no signs of an attractor that would preserve diversity.
However, a key difference between this experiment and that with the prey is that
the time required for all but one of the predator species to die out is significantly
longer than with the prey species. Figure 8 shows sample simulation runs with two
(top panel) and three (bottom panel) predator species, differentiated only by color,
and at a nutrient level of 0.20.
There is, of course, an attractor in terms of the total number of predators (the
sum of both predator populations, or the sum of all three populations) so that when
the population of one predator population dwindles, the other one (or other two)
compensate. In the top panel of Figure 8 especially, one can see that the variations
in population numbers is largely constrained to being between species.
As stated previously, the additional fish species generally died out within 1000
simulation time steps. As noted, for the predator population this process is signif-
icantly longer. The top panel (two predator populations) and bottom panel (three
predator populations) of Figure 8 illustrate two typical simulation runs of 100,000
time steps each. 100,000 time steps represents two orders of magnitude longer than
the similar results for the prey populations. We assume that this difference in re-
quired duration is an artifact of the less-realistic reproduction model inherent in
the predator agents; i.e., unlike the fish, these simulated agents spawn asexually.
As such, the predators are less dependent on spatial clustering, for example, and
are able to preserve diversity longer. Although it is unlikely, if there are mecha-
nistic reasons for this long persistence that are not simulation artifacts, it could
help us to understand the paradox of the plankton. For example in this simulation,
100,000 time steps represents ~150 predator lifetimes, which is generally well outside
a time-scale of interest.
Marine-Ecosystem
Fig. 8. Two illustrative simulation runs for Gause’s Law experiments with predator populations.
The top panel shows results with two predator species and the bottom panel represents the same
experiment with three predator populations.
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Emergent features in a general food web

Article in Advances in Complex Systems · December 2013

Ted Carmichael Mirsad Hadzikadic

SEE PROFILE SEE PROFILE

EMERGENT FEATURES IN A GENERAL FOOD WEB

Received (received date)

2 Carmichael and Hadzikadic

Emergent Features in a Food Web Simulation 3

tractable. In order to address certain limitations of purely mathematical models

2. Materials and Methods

4 Carmichael and Hadzikadic

Emergent Features in a Food Web Simulation 5

6 Carmichael and Hadzikadic

2.3. Modeling Methodology

Emergent Features in a Food Web Simulation 7

8 Carmichael and Hadzikadic

Emergent Features in a Food Web Simulation 9

10 Carmichael and Hadzikadic

Table 1. Predator and fish consumption levels.

Enrichment Level Predator Consumption Rate Fish Consumption Rate

Emergent Features in a Food Web Simulation 11

3.2. The Paradox of Enrichment

12 Carmichael and Hadzikadic

Table 2. Consumption rates, species additions, and species re-

Note: Fish and predator consumption rates (taken from Table

Emergent Features in a Food Web Simulation 13

3.3. The Competitive Exclusion Principle

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Emergent Features in a Food Web Simulation 15

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Emergent Features in a Food Web Simulation 17

4.1. Conclusions and Future Work

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Emergent Features in a Food Web Simulation 19

Based Approach, Marine and Coastal Fisheries: Dynamics, Management, and

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Emergent Features in a Food Web Simulation 21

tems in Ecological Time, Science 171 (1971) 385–387.

Appendix A. Stepped Pattern of Biomass Accrual - Extended

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Appendix B. Competitive Exclusion Principle - Extended Report

At initialization, fish were randomly assigned to ”red” or ”green.” The sim-

Emergent Features in a Food Web Simulation 23

24 Carmichael and Hadzikadic

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