Summary

Coral reefs are fascinating ocean habitats which harbor an incredible diver-
sity of marine life. The Lucero reef in the northwestern Philippines recently
experienced a severe drop in its biodiversity and health, roughly coinciding
with the increase in milkfish farming in Bolinao, Pangasinan. We establish
a mathematical model to represent the Lucero reef ecosystem and to aid in
predicting an economically viable solution that provides for the recovery
of the reef.
We reduce the ecosystem to a manageable complexity by modeling gen-
eralized trophic levels instead of individual species. In doing so, we limit
the precision of our model, but are still able to predict trends based on
pertubations to steady state conditions. Our model is based on the Lotka-
Volterra equations, diverging as necessary to accommodate the unique fea-
tures of the reef ecosystem. The model indicates that reducing milkfish
farming and increasing the harvest of algae and other primary producers
such as seaweed will benefit the reef ecosystem by limiting algae over-
growth. Moreover, previous research has shown that seaweed maricul-
ture is a satisfying economic solution. Thus we recommend a reduction in
milkfish harvesting to be compensated by an increase in algae harvesting.
This produces a compromise between the economic and environmental de-
mands on the system.
 Legalize (Sea)weed
ICM Contest Question C

Team # 5201

February 9, 2009

Contents
1 Introduction 2

2 Background 4
2.1 Selected Components of the Reef Ecosystem . . . . . . . . . 4
2.1.1 Coral . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2.1.2 Plankton . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2.1.3 Nutrients in the Environment . . . . . . . . . . . . . . 5
2.1.4 Milkfish . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2.1.5 Herbivorous fish . . . . . . . . . . . . . . . . . . . . . 5
2.1.6 Crustaceans . . . . . . . . . . . . . . . . . . . . . . . . 6
2.1.7 Echinoderms . . . . . . . . . . . . . . . . . . . . . . . 6
2.1.8 Molluscs . . . . . . . . . . . . . . . . . . . . . . . . . . 6
2.1.9 Algae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
2.2 Assessing Water Quality . . . . . . . . . . . . . . . . . . . . . 7

3 Model 7
3.1 Goals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
3.2 Existing Models . . . . . . . . . . . . . . . . . . . . . . . . . . 7
3.3 Assumptions . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
3.4 Our Models . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
3.4.1 Natural Ecosystem . . . . . . . . . . . . . . . . . . . . 9
3.4.2 Milkfish Monoculture Ecosystem . . . . . . . . . . . . 11
3.5 Model Limitations . . . . . . . . . . . . . . . . . . . . . . . . . 12

4 Integrated Multi-trophic Aquaculture Design 13

Page 1 of 24
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5 Results 13
5.1 Natural Ecosystem . . . . . . . . . . . . . . . . . . . . . . . . 13
5.2 Milkfish Monoculture . . . . . . . . . . . . . . . . . . . . . . . 13
5.3 Milkfish Mariculture . . . . . . . . . . . . . . . . . . . . . . . 14
5.4 Fisheries Management Design . . . . . . . . . . . . . . . . . . 17
5.4.1 Reduction of Milkfish Farming . . . . . . . . . . . . . 17
5.4.2 Integrating Seaweed Aquaculture . . . . . . . . . . . 17

6 Future Work 19

7 Conclusion 19

8 Appendix 20
8.1 Call to Action: Recommendations for Conservation Manage-
ment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
8.1.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . 20
8.1.2 Results of the Lucero Reef Model . . . . . . . . . . . . 21
8.1.3 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . 21

1 Introduction
Although coral reefs only cover about 0.1% of the ocean surface, they are
home to nearly one-third of all ocean fish species. They represent a micro-
cosm of biodiversity with profound implications for the health of the ocean,
yet over 20% of the world’s coral reefs have been destroyed due to human
activity and show no promise of recovery.[7] This is a sobering fact for the
state of our natural world, but it has immediate economic consequences as
well. It is estimated that healthy reefs proivde as much as $350 billion per
year in goods and services.[7] The combination of economic and environ-
mental implications incentives a strong mathematical model which can be
used to model the health of coral reefs and hopefully guide conservation
policy.
One reef which has been studied extensively in the literature is the
Lucero Reef off the coast of Santiago Island in the Philippines. The Lucero
Reef encapsulates many aspects of reef conservation. Historically it has
been a good model of tropical biodiversity, with a mean biodiversity in-
dex of H 0 = 2.60.[1] However, in 1995 milkfish harvesting became a new,
productive source of food and money and by 1998 there were an estimated
11,900 tons of milkfish farmed annually.[13] The milkfish farms have lead
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directly to increased nutrient levels in the reef area.[19] High levels of nu-
trients such as carbon, nitrogen, and phosphorous compounds have been
linked to algal blooms, which negatively impact coral life.[25], [19] As such,
these milkfish farms have a negative impact on the reef ecosystem, disrupt-
ing the delicate balance of biodiversity and threatening its long-term sur-
vival.
In creating conservation policy surrounding this reef ecosystem, it is
important to balance environmental and economic aspects of the issue. One
cannot ignore the commercial advantages of milkfish farming, both as a
source of food and income to island inhabitants, but taken to an extreme
milkfish farming can disrupt other local fishing industries if the coral reef
ecosystem collapses. Hence it is imperative both for environmental and
economic reasons to arrive at a sustainable solution for the island’s needs.
Aquaculture is the practice of designing controlled populations for fish
farming. Integrated multi-trophic aquaculture (IMTA) is a more recent ap-
proach in Western aquaculture that attempts to create a sustainable ecosys-
tem. The right balance of species provide resource recycling that preserves
a greater fish diversity, where the wastes of one become the resources of an-
other. This contrasts monocultures, high-densities of a single fish species
that propagate in regions where one species has become financially suc-
cessful, which are prone to deterioration through environmental degrada-
tion and disease outbreaks.[17] Many IMTA projects have begun through-
out the world in recent years, though we found no particular projects for an
ecosystem containing coral reefs. Our challenge is to incorporate the IMTA
framework specifically into the Bolinao region of Pangasinan, Philippines,
keeping the overarching goal of preserving the coral reef.
We began by creating a model of the Lucero Reef before the commercial-
ization of milkfish farming. The model is based off of the Lotka-Volterra
equation, using trophic levels to represent the various species in a reef
ecosystem. We then extended this model to apply to the milkfish farm-
ing pens, and obtained a high steady state value of nutrients and algae.
These values are detrimental to the coral, so we used our model to predict
changes that could be made to the mariculture ecosystem to reduce these
levels.
Team # 5201 Page 4 of 24

2 Background
2.1 Selected Components of the Reef Ecosystem
The reef ecosystem is intractably complex; modeling every component of
the ecosystem with good precision would provide ground-breaking insight
into the functional relationships within the ecosystem, but it would be ex-
ceedingly difficult. Realistically, to come to a conclusion in a reasonable
amount of time, we must reduce the ecosystem to relatively few general-
ized components and relationships. To reveal some of the intricacy of the
reef ecosystem and to illustrate its basic structure and dynamics, we de-
scribe its more prevalent components.

2.1.1 Coral
Most corals are colonial organisms consisting of hundreds of thousands
of individual polyps.[34] The reefs formed by some corals contribute sig-
nificantly to biodiversity by partitioning the environment and providing
for herbivores.[32] Reef-building corals often contain photosynthetic algae
called zooxanthellae, which offer a crucial mutualistic relationship to the
corals. While the coral protects the zooxanthellae and provides it with the
nutrients it requires for photosynthesis, the zooxanthellae removes wastes
from the coral and provides it with vital nutrients - oxygen, glucose, glyc-
erol, and amino acids. Because of this relationship between the coral and
zooxanthellae, coral tend to respond to the environment like plants, thriv-
ing in clear water that is easily permeable by sunlight.[36] Furthermore,
coral primarily exist in nutrient-poor water, as nutrient-rich water pro-
motes overgrowth of algae which can easily choke the coral. The removal
of the zooxanthellae, due to inadequate environmental conditions or other-
wise, is known as coral bleaching, and prolonged bleaching can cause the
coral to die from malnutrition.[35]

2.1.2 Plankton
Plankton are drifting organisms in oceans and other water environments.
Abundance and distribution of plankton depend heavily on nutrient con-
centration and other environmental conditions. Plankton are divided into
three functional categories: Phytoplankton, zooplankton, and bacterioplank-
ton. [30] Phytoplankton, a subset of algae, are microscopic organisms that
rely on photosynthesis for sustenance; as such they are primary produc-
ers, converting simple molecules and energy from sunlight into complex
Team # 5201 Page 5 of 24

organic molecules that are used by other organisms in the ecosystem.[29]

Zooplankton are small animals, including some crustaceans and larval stages
of larger animals like fish, that feed primarily on phytoplankton. Bacterio-
plankton are microorganisms including bacteria and archaea. [30]

2.1.3 Nutrients in the Environment

Reef nutrients take several forms, including sunlight and chemical com-
pounds such as ammonium, nitrite, nitrate, phosphate, and silicate.[31]
Such nutrients are the primary sources of growth for algae and other pri-
mary producers, so nutrient-rich waters promote overgrowth of algae and
make a poor environment for coral reefs. Another significant form of nutri-
ent found in the water is detritus, organic material that includes fragments
of dead organisms and feces.[10] Detritus is an important source of food
for much of the life in a marine ecosystem, including crustaceans and her-
bivorous fish.

2.1.4 Milkfish
Milkfish, or Chanos chanos, are a species of predatory fish found primarily in
the Indo-Pacific region along continental shelves and islands.[6] Their diet
consists primarily of small invertebrates, cyanobacteria, and soft algae.[6]
The average milkfish weighs roughly 1.5 kg[5], and in captivity is reported
to excrete 83 grams of waste per fish per day, which is composed of 11%
Carbon, 0.4% Nitrogen, and 0.6% Phosphorus. Based on this data, the
fish must consume about 30 kg per year. However, numbers reported by
Homer et al. (2002) on feeding rates indicated that milkfish consume only
1.2 kg per year. One report indicates that other omnivorous fish consume
roughly 30 kg per year per kg of biomass, which translates into roughly
45 kg per fish per year.[16] This number is on par with the estimate we ar-
rived at based on the weight of excrement, so we will use that estimate in
our model. Discrepancies in the paper by Homer, et al. are likely caused
by the fact that the reported consumption rate was based on supplimental
feed provided by milkfish farmers and certainly did not take into account
other sources of food.

2.1.5 Herbivorous fish

Herbivorous fish in reef habitats feed primarily on algae, corals, and phyto-
plankton. Besides this direct interaction with the coral habitat, they also af-
Team # 5201 Page 6 of 24

fect the environment indirectly by recycling nutrients through feeding and

defection.[32] We chose to study the representative species Scarus ghobban,
a member of the family Scaridae, more commonly known as Parrotfish.[27]
Parrotfish play an important role in the health of reef ecosystems, eating
the turf algae which would otherwise choke out the coral. They consume
approximately 109.1 grams of food per day, which translates into roughly
103 grams of turf algae per day.[27] Parrotfish have also been reported to
excrete 12 grams of waste per day.[27]

2.1.6 Crustaceans
The subphylum Crustacea is a large, diverse group of invertebrates. The
most well-known, and also the largest, crustaceans include crabs, lobsters,
and shrimp, However, there are many other, smaller varieties unknown in
popular culture.[8] Crabs, for example, are generally omnivorous, feeding
on algae, plankton, detritus, and small invertebrates. Smaller crustaceans,
like barnacles, feed primarily on plankton and detritus.[2]

2.1.7 Echinoderms
Echinoderms are a group of marine animals that live on the sea floor. Their
feeding habits vary significantly by species; echinoderms can be suspen-
sion feeders, herbivores, detritivores, and predators. The most common
groups in the Lucero reef, sea urchins and sea cucumbers, feed primarily
by grazing on algae. [28] Sea urchins therefore serve an important role to
coral health by keeping algae growth in check.

2.1.8 Molluscs
Molluscs, particularly bivalves, such as clams, fill an important niche in
reef ecosystems. Most bivalves feed by using large gills to capture nutrient
particles directly from the ocean water.[3] Bivalves thus reduce the nutri-
ent concentration in the water, thereby limiting the growth of algae and
bacteria. While this in turn limits the growth of herbivores who feed on
the algae, it positively affects the health of the reef by keeping algae from
overgrowing the coral.

2.1.9 Algae
Algae are a very broad group of photosynthetic, multicellular and unicel-
lular organisms. Most algae are primary producers, meaning they are an
Team # 5201 Page 7 of 24

essential part of the food chain that converts light and chemicals from their
environment into biomass. A representative species for our model is Ge-
lidium pusillum, a red algae[21] common in the province of Pangasinan.[33]
Gelidium pusillum forms algal turfs,[22] which are apt to overgrow and choke
coral. Algal turfs also trap detritus and form sites for bacterial growth,
thereby providing food for detritivores and some herbivores.[32]

2.2 Assessing Water Quality

Algal blooms have been directly linked to coral death in several studies
dating from the mid-1970’s.[25] As such, algal biomass in a reef ecosystem
is strongly correlated to the health of the coral and hence the health of the
entire system. Although there is debate in the literature,[24] algal blooms
have been linked by several studies to a combination of increased nutrients
(such as nitrates and phosphates) in the water and a decrease in grazing
from herbivorous species.[25],[26] Because nutrients impact algae growth
and thus indirectly impact the health of coral, it is important to model this
relationship. In our model, nutrient levels will be measured by a water
quality score, which takes into account the nitrates and phosphates linked
to algal blooms.

3 Model
3.1 Goals
We want to create a model that incorporates the entire foodweb of the Boli-
nao coral reef ecosystem. Data we wish to retain from the model includes
water quality levels, amount of fish harvesting, and steady state levels of
fish population. This values are ultimately used to evaluate suggestions for
an integrated multi-trophic aquaculture for the Bolinao region.

3.2 Existing Models

There is a long history of models on the lower part of the food chain that
model the interaction of phytoplankton, zooplankton, and nutrients. Other
models, particularly those used in fishery management decisions, rely on
survey data to model fish populations at a higher trophic level while ignor-
ing the bottom.
Furthermore, ecology modeling comprises of two distinct approaches,
state variable and individual-based modeling. State variables encapsu-
Team # 5201 Page 8 of 24

late the many facets of a species population into a single state, typically
the biomass of the species. This disregards important distinctions among
the individuals. In some cases, aggregation among the individuals was
found to correlate to similar patterns in a state variable model. However,
individual-based modeling is more often used on more narrow scales than
which we are concerned for this problem,[23] so we will focus on a state
variable model.
The integration of multiple trophic levels is a difficult modeling prob-
lem still today that requires a great deal of research. [18] A 2003 survey of
ecosystem models found that ECOPATH was that only model that used a
multispecies approach:

Despite broad discussion of the need to consider multispecies

issues in marine conservation and fisheries management we know
of only one model that focuses explicitly on multispecies as-
pects of marine reserves. [12]

Population dynamics are based on a conservation of mass principle,[9]

where the change in biomass of a species over time is based on growth
(sources of mass) or loss (sinks of mass). Loss can come from being eaten
by another organism or natural death. To conserve mass, some of the loss
of one state’s mass becomes a sink for another’s. The remaining mass ex-
creted as nutrients or other particles. Either growth or loss can occur from
migration. The Population Law of Mass Action describes these rates of
changing mass. It states that the rate of change of the mass of two interact-
ing species is proportional to the product of their masses. This principle is
used in modeling predator-prey relations in the Lotka-Volterra model,

x 0 = (− a + by) x = − ax + bxyy0 = (c − kx )y = cy − kxy

where x is the predator mass and y is the prey mass. The bxy and −kxy
terms represent the predation, a source of growth for the predators but a
sink for the prey. To close the model, − ax represents the natural death of
the predator (i.e. in isolation of the prey), and cy represents the natural
growth of the prey. [4]

3.3 Assumptions
The life-cycle of a marine organism involves distinct phases. However, we
assume that all biomass of a species has the same consumption pattern.
Team # 5201 Page 9 of 24

This pattern is determined as a rough average of consumption patterns

over all stages of life, weighted by biomass.
Nutrient uptake by phytoplankton is a function of both light and tem-
perature since it is dependent on the amount of photosynthesis that the
organism can perform.[9] We assume that nutrient uptake can be approxi-
mated as a temperature-invariant and light-invariant function.
Our models do not incorporate variations due to seasonal cycles. For
instance, the vertical mixing of ocean layers occurs in greater proportion in
winter than summer, which effects the flux of organisms and particles.[18]
They also do not take into account the acidity or temperature of the water,
bacteria, or virus levels.
Additionally, because it is not feasible to model every single species in a
very complex ecosystem, we have chosen to represent generalized trophic
groups instead of individualized species. Any attempts to use the specific
grazing patterns of any one species as representative for the entire trophic
group lead to nonsensical results because it assumes a closed system when
in reality there are many other species and factors at play. For example,
if we chose precise initial conditions for our primary producers off of the
absolute biomass of algae, our ecosystem would be underfed because it ne-
glects other primary producers such as seagrass and seaweed. Instead, we
chose to look at rough production and consumption ratios for each trophic
group and then force the grazing coefficients to achieve a steady state. Al-
though this does not precisely reflect reality, it gives sensical results and we
claim that the grazing coefficients represent a complicated mass balancing
that occurs behind the scenes in the way we calculated them.

3.4 Our Models

3.4.1 Natural Ecosystem
We began by creating a simplified model of the reef ecosystem based off
of the Lotka-Volterra equation. Algae compete for nutrients, and are eaten
by herbivores which represent trophic level III (which would include crus-
taceans and echinoderms). It is widely accepted and several sources con-
firm that only about 10% of the energy at any trophic level moves up to the
next trophic level, which we represent in our model by an efficiency coeffi-
cient e.[16] The herbivores are eaten by predatory fish, and finally there is
an extinction term that represents the rate at which the algae, herbivores,
and predators die and are returned to the nutrient sink. Because our system
looses energy at each trophic level (mirroring the 90% consumption of en-
Team # 5201 Page 10 of 24

ergy in biological systems for basic life processes), we introduced a constant

term α which represents energy flowing into the system and augments the
nutrient term consumed by the algae. This term can be conceptualized as a
measure of the sunlight adding energy to the ecosystem. Below is a math-
ematical representation of our model:

N 0 = dH + dA + dM + 0.9 ∗ gm MZ − ga N A
A0 = ga ( N + α) A − gh AH − dA
H 0 = egh AH − egm HM − dH
M0 = egm HM − dM

Table 1: Initial Conditions

Symbol Name Initial Value

N Nutrients 0.032
A Algae 0.3
H Herbivores 50
M Predators 0.92

We chose initial conditions based on estimations of biomass consump-

tion and production per year for each class of animal.[16] The algae is
represented by phytoplankton, the herbivores are represented in general
by Parrotfish, and the predators are represented by milkfish.1 Nutrient
levels were chosen to be a low steady state value based on data in the
literature.[15] We assumed that these numbers represent a stable equilib-
rium, and hence solved the set of steady-state equations:

0 = N 0 = dH + dA + dM + 0.9 ∗ gm MZ − ga N A
0 = A0 = ga ( N + α) A − gh AH − dA
0 = H 0 = egh AH − gm HM − dH (1)
0
0 = M = egm HM − dM
1 In reality, milkfish are not solely predators. However for the purposes of this model we

are assuming that they only eat herbivores

Team # 5201 Page 11 of 24

The death rate and efficiency coefficients were set to

d = .125
e = .1

as a way to calibrate the model. The efficiency parameter mirrors the fact
that only 10% of energy is passed up each trophic level, and the death rate
was set heuristically assuming that the biomass of each population is recy-
cled every eight years. The result of these calculations is the set of parame-
ters shown in table 2 which solve the steady state equations (1).

Table 2: Simplified Model Steady State Parameters

Symbol Name Value

α Resource Input 0.2865
e Energy Efficiency 0.1
d Death Rate 0.125
ga Algae Grazing Rate 774.7
gz Herbivore Grazing Rate 4.93
gh Predator Grazing Rate 0.025

When perturbed from the steady state, this model behaves as expected.
An increase in predator biomass causes a decrease in herbivore biomass
and hence an increase in algae biomass. Similarly, an decrease in preda-
tor has the reverse effect, and an increase in herbivore biomass results in
a decrease of algae biomass and increase in predator biomass. However,
over several generations these transient responses even out and return to
close to the original steady state. This corresponds to what we see in the
natural world. Within reason, perturbations from natural steady state val-
ues does not cause the entire world ecosystem to collapse; instead, there is
a transient flux until the system can find a new equilibrium. Our model,
although simplified, also has this characteristic.

3.4.2 Milkfish Monoculture Ecosystem

In many ways the milkfish mariculture system in the Philippines alters the
way in which the reef ecosystem operates. We chose to model the ecosys-
tem inside the fish farm pens where there is a high density of predator (ie.,
milkfish) in order to discover what effects fish farming has on the steady
state values of algae (which is our indicator of coral survival). Using the
Team # 5201 Page 12 of 24

simplified ecosystem model above, we modified some of the conditions to

reflect the nature of the milkfish monoculture ecosystem.
If we were to use the exact same model, the extremely high density of
milkfish in the pens would overwhelm the herbivore population and hence
lead to a near exponential growth of algae. Additionally, because the milk-
fish consume the herbivore population so quickly, it would lead to a rapid
decline in milkfish population after an initial peak. However, these effects
result from some simple assumptions that change in a mariculture ecosys-
tem. First of all, the milkfish are being fed and harvested so that the total
population remains almost constant. Hence, we can assume that M0 = 0.
Another difference between the mariculture and natural ecosystem is that
in the mariculture environment, milkfish derive most of their diet from the
food given to them by the farmers. We estimated that farmed milkfish only
eat one tenth as much from the herbivore group as they do in a natural
environment. Finally, because the milkfish are being farmed, their dead
biomass does not return to the nutrient compartment. These changed as-
sumptions resulted in the following set of differential equations:

N 0 = dH + dA + 0.9 ∗ gm MZ − ga N A
A0 = ga ( N + α) A − gh AH − dA
H 0 = egh AH − 0.1 ∗ gm HM − dH
M0 = 0

These equations will be solved numerically given an elevated milkfish

population to study the effects of fish farming on the reef environment.

3.5 Model Limitations

The simplification we made by representing different species by their gen-
eral trophic level is a significant limitation to the model. It does not take
into account the varied grazing patterns of different organisms, and does
not have very high resolution to look at the ecosystem on a per species
level. Additionally, it neglects the complexity of the detritus cycle by as-
suming that nutrients get converted into algae biomass with perfect effi-
ciency. This leaves out whole classes of organisms such as molluscs which
can feed off of dendrite. It also ignores the fact that very few species are
truly either carnivores or herbivores. In reality, milkfish (which were the
focus of our model) feed off of algae and detritus as well as off of small
fish, floating fish eggs, and small invertebrates. However, our model tries
Team # 5201 Page 13 of 24

to capture the general flow of energy between trophic levels, and even af-
ter the simplification can give some meaningful physical results. However,
any interpretation from our data must be taken in the context of the model;
our results are not meant to show conclusively that any one outcome will
occur given a certain set of input. Instead, it is meant to give a general trend
for different perturbations to the reef ecosystem which may not be readily
obvious upon inspection. Further development is needed for more precise
quantitative measures.

4 Integrated Multi-trophic Aquaculture Design

Developing a sustainable ecosystem for the Bolinao coral reef involves a
delicate balance. Excess nutrients allow the growth of algae, which com-
pete with coral for sunlight used in photosynthesis by covering surface area
of the ocean.
We agreed with the several sources proclaiming the importance of build-
ing a resilient coral reef ecosystem. By this, we mean the ability of the reef
to withstand unforeseen major changes.

Reef ecosystems seem to shift between alternative stable states,

rather than responding in a smooth way to changing conditions.
The shift to algae in Caribbean reefs is the result of a combina-
tion of factors that make the system vulnerable to events that
trigger the actual shift.[14]

As an example of resilience, the large number of sea urchins allowed

the Caribbean reef to recover from a 1981 hurricane that destroyed much of
it. it soon after suddenly was severely hindered by brown algae.

5 Results
5.1 Natural Ecosystem
We calibrated our model to a steady state representing the species popula-
tions in the coral reef ecosystem prior to milkfish farming in Bolinao.

5.2 Milkfish Monoculture

We next modeled the ecosystem of the fish pens after milkfish farming was
introduced to Bolinao. To simplify the model, milkfish were set to the pop-
Team # 5201 Page 14 of 24

Figure 1: With no herbivorous species, our model predicts algae growing

without bound.

ulation described by [13]2 and the herbivore species were reduced to zero.
The nutrient levels greatly exceed the threshold determined by [25]. Com-
bined with a zero population of herbivores, this fact leads to the uncon-
trolled growth of algae seen in the above figure. Clearly this result is not
physical. Algae would eventually reach a the carrying capacity of the envi-
ronment, and it is also unlikely that the herbivore population would ever
be zero. However, these limitations of our model do not detract from the
fact that a low herbivore population and high milkfish population would
lead to dangerously high levels of algae which would threaten the coral.

5.3 Milkfish Mariculture

We adjusted the fish pen ecosystem model by setting the herbivore popula-
tion to 500 g/m2 such that we were able to get a nutrient level correspond-
ing to the current levels as reported by [20]. This population level compares
to 247 g/m2 , the actual populations in Bolinao.[16] The herbivore popula-
tion would need to increase to compensate for the algae blooms caused by
waste nutrients from the milkfish.
2 We calculated a milkfish biomass of 10 kg/m2 inside the pens.
Team # 5201 Page 15 of 24

Figure 2: Species biomass in present Bolinao fish pen conditions over 10

year span.

Figure 3: Algae biomass in Bolinao over 31 weeks. The biomass spikes to

about 150 g/m2 about a week after the milkfish are added.
Team # 5201 Page 16 of 24

Figure 4: Nutrient levels in present Bolinao fish pen conditions over 10 year
span. (Steady state value is approximately 2.5 g/m2 )

Figure 5: Herbivore biomass (g/m2 ) present in Bolinao fish pen conditions

over 10 year span.
Team # 5201 Page 17 of 24

5.4 Fisheries Management Design

5.4.1 Reduction of Milkfish Farming
Since large levels of algae threaten the health of reefs, we first investigated
a simple way to reduce the amount of algae. Limiting the mass of milkfish
by half and then to 10% decreased the levels of algae from about 20 g / m2
to near 10 g / m2 and 2 g / m2 , respectively.

Figure 6: Algae levels decreased proportional to the reduction in milkfish

farming

5.4.2 Integrating Seaweed Aquaculture

An alternate idea to simply attempting to decrease milkfish levels is to in-
corporate the idea of a integrated multi-trophic aquaculture by introduc-
ing seaweed to the fish pens. This keeps the algae from the possibility of
damaging the coral and also allows the profitable harvest of milkfish. Our
model predicts that we can harvest 2 kg/m2 of algae per year without de-
stroying the ecosystem. At a calculated rate of $1.40 per kilogram of algae,
this comes out to roughly $3,000 per pen per year.[11]
As indicated in the final figure, this model unfortunately predicts a de-
crease in the herbivore life in the fish pens. However, a healthy coral reef
due to lower nutriet levels may still allow the herbivores to thrive there.
Team # 5201 Page 18 of 24

Figure 7: After introducing seaweed harvesting, nutrient levels decreased

to an acceptable level.

Figure 8: An equilibrium state of algae is still maintained after harvesting

seaweed from the fish pens.
Team # 5201 Page 19 of 24

Figure 9: Herbivore life in the fish pens is reduced to a low level.

6 Future Work
Our model provides only a glimpse in the complex ecosystem represented
in Bolinao. A more accurate model, incorporating the variety of omissions
characterized previously, would give a better representation of the changes
that would occur when testing the possibilities for a new polyculture. Ad-
ditional data to improve the accuracy of parameters of the model would
provide similar improvements.
The great diversity of sea life means that many solutions may poten-
tially exist to a integrated multi-trophic aquaculture in Bolinao. This is an
area of active research, and keeping an eye on the developments in similar
fisheries, particularly in coral reefs, may provide some additional insight
into designing a management that provides continued harvesting value
through a sustainable ecosystem.

7 Conclusion
Milkfish farming destroyed the once diverse aquatic life and beautiful coral
of Bolinao. But the technology of integrated multi-trophic aquaculture can
change that, allowing the coral to rebuild. Many fisheries are now being
managed with a sustainable recycling of wastes while still supporting an
acceptable level of harvesting. Our model demonstrates that this result is
a possibility in Bolinao. Reducing the penned milkfish by a half and in-
Team # 5201 Page 20 of 24

troducing other forms of harvestable herbivores leads to a lower steady

state value of algae. Combined with the harvesting of algae this can lead
to a viable economic and environmental solution. Our models showed that
perturbations to the steady state (through constant harvesting terms) does
little to affect the overall stability, leading to the conclusion that high lev-
els of harvesting can be sustained. We believe a conservation policy built
around this model has a decent chance at reversing the trends seen today
in the Bolinao reef system.

8 Appendix
8.1 Call to Action: Recommendations for Conservation Manage-
ment
8.1.1 Introduction
Conservation management is an important issue facing many nations as
they try to balance the economic and food needs of their country with the
needs of natural ecosystems. Nowhere is this balance more pronounced
than in coastal reef ecosystems, which house over one third of all ocean
fish species and a vast amount of the biodiversity found in ocean envi-
ronemnts. The incredible complexity of these ecosystems also makes them
incredibly fragile; as such, conservation management becomes especially
important. We realize fully the difficult challenges posed by creating sta-
ble, viable ecosystems which also produce harvestable product. Our team
of researchers has created a mathematical model to assist with these chal-
lenges, providing a way to visualize the impact of different changes to the
system. In modeling the Lucero Reef system on the coast of Pangasinan,
Philippines, we found that milkfish farming could be sustainable given a
controlled number of fish and a polyculture of other organisms. We rec-
ommend cutting the number of farmed milkfish in half and making up for
lost profits through harvesting algae. This would not only keep algae at
acceptable levels for coral survival, but provide a viable economic model
which includes food and income for island inhabitants.
In our readings of the literature, we found a strong correlation between
algae levels and coral health. If algae becomes too pervasive, they compete
with the coral for vital resources and effectively ”choke out” the coral. Be-
cause coral is such a vital piece of the reef ecosystem, coral death would ul-
timately lead to the ecosystem’s collapse. This is an outcome any conserva-
tion management program should be designed to avoid. Below we present
Team # 5201 Page 21 of 24

specific results from our model of the Lucero Reef to justify our claims. We
then provide an economic assessment, demonstrating a strengthening of
the Bolinao mariculture industry under our proposed plan.

8.1.2 Results of the Lucero Reef Model

We first modeled the Lucero Reef ecosystem before the introduction of
milkfish farming. Our model demonstrated stable steady state values cen-
tered on estimates of relative biomasses of three main trophic levels, namely,
the primary producers, herbivores, and predators. Perturbations to this
system behaved in the way we expected, quickly returning to the steady
state values. This demonstrates the robustness of the reef ecosystem, mod-
eling in a simple way a complex polyculture of organisms which keep each
other in balance.
The model was then extended to apply to a mariculture ecosystem. Cur-
rently, milkfish farming is common along the outer edges of the reef. In the
milkfish pens, the steady state of the ecosystem is disrupted. High levels of
nutrients from the fish food and excrement lead to high levels of algae. We
found that reducing the milkfish from 10,000 g/m2 to 5,000 g/m2 in com-
bination with introducing new herbivores would cut the algae to 10 g/m2 .
Although this is still much higher than an acceptable value, harvesting the
algae could bring the number down to an environmentally friendly range.
Algae is reported to have brought in $6.2 billion in 2008, so harvesting this
aglae for economic profit is a viable commercial solution to an environmen-
tal problem. Our model predicts that we can harvest 2 kg/m2 of algae per
year without destroying the ecosystem. At a calculated rate of $1.40 per
kilogram of algae, this comes out to roughly $3,000 per pen per year.

8.1.3 Conclusion
We suggest implementing these recommendations quickly to reverse the
detrimental trends present in the Bolinao reef system. With a conscious
effort, conservation management can produce a vibrant economic and en-
vironmental solution to this problem.

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