Reviews in Aquaculture (2010) 2, 73–85 doi: 10.1111/j.1753-5131.2010.01023.

Shrimp polyculture: a potentially profitable, sustainable,

but uncommon aquacultural practice
Marcel Martı́nez-Porchas1, Luis R. Martı́nez-Córdova2, Marco A. Porchas-Cornejo3 and
José A. López-Elı́as2
1 Department of Technology of Foods of Animal Origin, Centro de Investigación en Alimentación y Desarrollo (CIAD), Km 0.6 Road to
La Victoria, ZC 83304, Hermosillo, Sonora, México
2 Department of Scientific and Technological Research of the University of Sonora, Universidad de Sonora, Luis Donaldo Colosio Blvd. w ⁄ n,
Colonia Centro, ZC 83000, Hermosillo, Sonora, México
3 Center of Bological Research, Centro de Investigaciones Biológicas del Noroeste (CIBNor), Unidad Sonora, Guaymas, Sonora, México

Correspondence
Marcel Martı́nez-Porchas, Centro de
Investigación en Alimentación y
Desarrollo, A.C. Km 0.6, Carretera a
La Victoria, Hermosillo, Sonora.
Email: marcelmp_6@hotmail.com;
marcel@ciad.mx
Received 27 August 2009; accepted
19 November 2009.
Abstract
Shrimp polyculture is not yet a common practice among farmers; however, this
activity represents an important alternative to solving and ⁄ or minimizing some
of the problems that shrimp aquaculture has faced in the past two decades
(environmental pollution, diseases and decreasing prices). In this context, many
benefits have been achieved with some polyculture practices. Several species
from diverse trophic levels have the potential to be co-cultured with shrimps.
A good knowledge of the species that are candidates for polyculture and an
adequately designed culture system are the most important points to consider
when co-culturing shrimp with other species. The present paper is a review of
the past, present and future of shrimp polyculture with other organisms.
Key words: aquaculture, co-culture, environmental pollution, shrimp farming, shrimp industry,
sustainable aquaculture.

and Agriculture Organization of the United Nations

Introduction
Aquaculture is an important worldwide activity, providing
a source of food and economical profit for humans
(Martı́nez-Córdova 1998). Hundreds of aquatic species of
fishes, crustaceans, mollusks and plants are farmed in
many countries and many others are being studied with
that purpose (FAO 2007).
Crustacean culture is, in general, one of the most profit-
able aquaculture activities because the generated products
have high commercial value, representing an annual eco-
nomic profit of US$14 361 million (FAO (Food and Agri-
culture Organization of the United Nations) 2007). In this
context, penaeid shrimps are one of the most important
aquaculture organisms, particularly ‘white’ or ‘crystalline’
shrimps such as Litopenaeus vannamei, Litopenaeus setiferus
or Litopenaeus stylirostris. Other penaeid shrimps that are
being successfully cultured are Penaeus monodon, Penaeus
semisulcatus, Marsopenaeus japonicus, Fenneropenaeus
chinensis, Fenneropenaeus indicus, Fenneropenaeus penicilla-
tus, Farfantepenaeus aztecus, Farfantepenaeus californiensis,
Farfantepenaeus paulensis and Farfantepenaeus duodarum
(Martı́nez-Córdova & Campaña-Torres 1999). The Food
calculated a total of 375 913 shrimp farms in 2000 all over
the world.
Despite the benefits and progress of penaeid shrimp
aquaculture, this activity faces several problems, including
disease (Morales-Covarrubias 2004), environmental pollu-
tion, particularly as a result of farm effluent (Sansanayuth
et al. 1996), and decreasing prices (FAO (Food and Agri-
culture Organization of the United Nations) 2007). Some
strategies, such as antibiotic administration (Bachère
2000), the use of probiotics (Vieira et al. 2007), recircula-
tion systems (Otoshi et al. 2003), bioremediation of efflu-
ents (Sangamaheswaran et al. 2001; Paniagua-Michel &
Garcı́a 2003) and polyculture with other species (Eldani
& Primavera 1981; Saelee 2002; Thu 2003; Martı́nez-
Córdova & Martı́nez-Porchas 2006), have been used to
minimize these problems.
According to Lanza-Espino et al. (1991, pp. 101–120),
polyculture consists of ‘adding one or more subordinate
species to the culture system of a main species’. In recent
years, research into crustacean polyculture has increased,
demonstrating in some cases that it is not only a profit-
able, but also a sustainable activity, both in freshwater

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M. Martı́nez-Porchas et al.
and marine systems (Irz & Mckenzie 2003; Rahaman
2006). Moreover, some authors have found that adding a
secondary or subordinate species improves the perfor-
mance of the main cultured organism (Gonzales-Corre
1988; Wang et al. 1998; Tian et al. 2001). For instance,
different species of mollusks (Crassostrea gigas and Chione
fructifraga) and fish (Oreochromis urolepis hornorum)
(Martı́nez-Córdova & Martı́nez-Porchas 2006; Tendencia
et al. 2006) have been co-cultured with penaeid shrimps,
improving the production parameters of the shrimps.
Unfortunately polyculture is not yet a common practice
in shrimp farms, in some cases because of the farmer’s
fear of losing the shrimp harvests to the possible intro-
duction of pathogen viruses or bacteria into the system,
and in other cases because a lack of knowledge of poly-
culture principles and practices. For example, in the state
of Sonora, which is the major producer of shrimp by
aquaculture in Mexico, with 70% of the total production
(approximately 100 000 tons year)1) (Gutierrez-Venegas
2006), a recent survey conducted by the authors
(M. Martı́nez-Porchas et al., unpubl. data, 2008) revealed
that more than 80% of the producers did not have infor-
mation about polyculture practices and none had consid-
ered using polyculture. In addition, Bunting (2008)
demonstrated that people involved with aquaculture were
pessimistic with regard to the financial returns from poly-
culture.
The present paper is a review of the benefits, strategies,
principles and practices of shrimp polyculture around the
world, with the aim of encouraging the scientific commu-
nity and shrimp producers to develop this important
activity.
Problems in the shrimp industry
In past years, the aquaculture industry, particularly
shrimp culture, has resulted in contamination of the envi-
ronment and the destruction of millions of hectares of
mangle forest in different countries (Naylor et al. 2000).
Other problems related to the aquaculture industry
include eutrophication and nutrification in discharge-
receiving ecosystems, which have generated explosive
growth of phytoplankton (sometimes toxic species), the
burring of benthic species, undesirable odors and the pro-
liferation of pathogenic microorganisms. Alterations to
the hydraulic patterns and landscape (affecting the tour-
ism industry) are some other situations attributable to
aquaculture (Martı́nez-Córdova et al. 2009).
In contrast, the price of shrimp has dropped abruptly
in recent years, although the price of supplemental feeds
has not shown the same tendency. This affects the eco-
nomical profitably of the shrimp industry and makes it
necessary to look for alternative strategies to optimize the
74
management of feed and feeding (Martinez-Córdova
2009). In this context, polyculture is one of the most
promising alternatives; in fact Lebel et al. (2009) pro-
posed that ‘polycylture practices’ and other strategies
would minimize the severe problems that the shrimp
industry is facing and generating in Mexico and Thailand.
Polyculture types
Polyculture is also referred to as multi-trophic aquacul-
ture, co-culture or simply integrated aquaculture (Bunting
2008). There are at least three general types of polycul-
ture: direct polyculture, cage-cum-pond polyculture and
sequential polyculture (Yi & Fitzsimmons 2004) (Fig. 1).
Direct polyculture refers to two or more species mixed
in the same pond or aquaculture unit without partition-
ing (i.e. it is possible that there will be physical contact
between the species). This type of polyculture does not
require significant extra economical or human effort in
investment because the secondary species are simply
added within the same space of the main species; thus,
there is no necessity for more space. However, in some
cases, extra aeration and an increase in the water-
exchange rate may be required. For example, Gonzales-
Corre (1988) stocked juveniles of the shrimp Pena-
eus monodon (PL 45) and after 35 days he added tilapia
fingerlings Oreochromis niloticus (2.01 g) and found that
the concentration of oxygen tended to decrease as the
density of tilapia increased.
Direct polyculture is the simplest type of co-culture
and can be carried out on almost any farm. However, this
activity is limited by species compatibility and the loading
capacity of the system. One of the most important con-
siderations should be the food requirements and the envi-
ronmental necessities of the candidate species that are to
be added to the culture of the shrimp (Pitt & Duy 2004;
Douglass et al. 2008). In addition, the density of organ-
isms within the system may be a limiting factor because
oxygen consumption increases as a function of biomass.
For instance, it is not possible to add a subordinate spe-
cies into an intensive or super-intensive shrimp culture
(Martı́nez-Córdova 1997) without supplemental aeration.
Cage-cum-pond polyculture is similar to direct poly-
culture because different species are cultured together in
the same tank or pond, but one (or some) of them are
enclosed within cages, limiting their space and physical
contact with the main organism (Yi & Fitzsimmons
2004). As with direct polyculture, no extra space is
needed and furthermore there is better control of the cul-
tured species. Obtaining the cages may require an extra
economical investment; however, this investment may not
be very high because some types of cages can be con-
structed from cheap materials such as polyester ⁄ polyvinyl
Reviews in Aquaculture (2010) 2, 73–85
ª 2010 Blackwell Publishing Asia Pty Ltd
 Shrimp polyculture, a sustainable practice

(a)

(b)

(c)

Figure 1 Polyculture types. (a) Direct polyculture, (b) cage-cum-pond polyculture and (c) sequential polyculture. Different organisms may be
cultured than those in the images. Images are sourced from: http://www.fishbase.org and http://www.teara.govt.nz/1966/T/Tuangi/Tuangi/
ENZ-Resources/Standard/1/en.

chloride and plastic net. Depending on the mesh screen

size, some cages may require continuous maintenance
because organisms and organic matter that restrict the Shrimp Shrimp Shrimp Shrimp
circulation of water and oxygen may cover the walls and pond pond pond pond
bottom of the cages. This activity is limited by the pond
space and the biomass loading capacity. Cages cannot be
used in regions with extreme temperatures because the
water surface where the cages are installed may reach
Weeds Weeds Weeds Tilapia
extreme temperatures and the organisms within the cages
pond pond pond pond
are not allowed to move toward the bottom. In contrast,
a cage cannot be placed at the bottom because during the
night the oxygen decreases to suboptimal or even lethal Figure 2 Integrated, multitrophic and recirculation shrimp polycul-
levels (Lanza-Espino 1998). ture system proposed by Ruenglertpanyakul et al. (2004).
Sequential polyculture implies an integrated aquaculture
system where the main and subordinate species are sepa- Unfortunately, these types of systems can require much
rated by different units and the shrimp effluents flow more economical investment, are time consuming and take
toward the culture units of the subordinate species up extra space. Chien and Tsai (1985, p. 430) argued that
(Fig. 2); in some of these systems the water is discharged when in a polyculture the ‘organisms have different physi-
to receiving ecosystems, whereas in others it is re- cal needs or have antagonistic (competitive or predator–
circulated (Chien & Tsai 1985). The secondary species prey) relationships, successful polyculture (within the same
thrive in the effluents from the ponds (or tanks) of the structure) becomes unlikely’, and that an alternative solu-
main species, feeding on non-consumed food, organic tion can be the use of ‘integrated pond culture’ systems.
matter and other nutrients. This process improves the In all three general types of polyculture there are two
quality of the discharged water so that it can be reused and subdivisions; the first is ‘monotrophic aquaculture’ where
decreases the environmental impact of the aquaculture. similar species (in terms of nutrition) are added to the

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ª 2010 Blackwell Publishing Asia Pty Ltd 75
M. Martı́nez-Porchas et al.
culture system, for example, two or three species of
shrimp (Martı́nez-Córdova & Peña-Messina 2005). The
other subdivision is ‘multitrophic polyculture’ where
organisms from different trophic levels are added to the
system (e.g. shrimp–fish–seaweed). Troell et al. (2009,
p. 3) asserted that a polyculture can ‘combine fed aqua-
culture species (e.g. finfish or shrimp), with inorganic
extractive aquaculture species (e.g. seaweeds) and organic
extractive species (e.g. suspension- and deposit-feeders)
cultivated in proximity’.
Polyculture benefits and disadvantages
Benefits
Many benefits have been achieved in shrimp polyculture
systems when using fish, bivalves and seaweed as subordi-
nate species, despite the fact that polyculture is not com-
mon and rarely researched. These benefits have been
achieved in all three polyculture systems (direct, cage-
cum-pond and integrated polyculture); the benefits
include the diminution of ecological impacts and
improvements in yield and water quality (Muangkeow
et al. 2007; Troell et al. 2009).
Polyculture can contribute to minimizing the environ-
mental impact of farm effluents, particularly those related
to nitrogenous wastes, which are further converted into
toxic metabolites; the main reason for this is that some
subordinate species can fed on and assimilate most of the
wastes generated from shrimp aquaculture. A higher effi-
ciency of nitrogen utilization has been observed in poly-
culture systems compared with monoculture systems
(Zhen-xiong et al. 2001), with a consequent decrease in
nitrogen excess, improvement in water quality and dimi-
nution of the environmental impact resulting from efflu-
ent discharges. Yokoyama et al. (2002, pp. 745–746)
asserted that ‘in polyculture systems, the wastes from
aquaculture are assimilated through the food web within
pond microcosm formed by the co-cultured organisms
and natural pond biota’. There is also evidence that the
diversity of species within a specific environment influ-
ences a variety of ecosystem processes including produc-
tivity, decomposition and nutrient cycling (Hooper et al.
2005; Balvanera et al. 2006; Douglass et al. 2008). Belton
and Little (2008) affirmed that shrimp culture has
severely affected the ecosystem and concluded that inte-
grated aquaculture practices, such as polyculture, are
good alternatives for reducing contamination.
Marine organisms such as sea cucumbers that live in
bottom soil can recycle nutrients and bioturbate sedi-
ments, limiting the proliferation of anaerobic bacteria and
the consequent formation of anoxic zones (Uthicke
1999). Nile tilapias and other fish can filter feed phyto-
plankton and organic wastes, which may eventually affect
76
the physiological status of the shrimp and also be a
potential contaminant in environments receiving farm
effluents (Tookwinas 2003; Turker et al. 2003). Oysters
and other bivalves have been shown to remove suspended
particles that are reported to have a negative effect on the
growth of shrimp and on the environment (Jones et al.
2001, 2002). Shpigel (2005) pointed out that the use of
suspension feeders such as bivalves in biofiltration
represents a cheap alternative for the biological
removal of nutrients from effluent waters. Bunting (2006)
constructed an integrated shrimp–shellfish–seaweed
polyculture system and observed that hairy cockle (Scaph-
arca inaequivalvis) and seaweed (Gracilaria spp.) reduced
the total ammonia nitrogen (TAN), total nitrogen (TN)
and total phosphorous (TP) concentrations from shrimp
effluents by 61, 72 and 71%, respectively. Seaweed has
been shown to remove nutrients from the effluents of
aquaculture farms (Nelson et al. 2001; Yarish et al. 2001;
Marinho-Soriano et al. 2002). Kaewsuralikhit (1994) and
Khidprasert (1995) found that red seaweeds (Gracilaria
salicornia and Gracilaria fisheri), green seaweed (Caulerpa
macrophysa) and brown seaweed (Sargassum polycystum)
were capable of effectively assimilating TAN, NO2 and
NO3 (>40%). All these reports clearly indicate that a
potential alternative to diminish the pollution associated
with aquaculture effluent is bioremediation throughout
seaweed species that are capable of assimilating and using
the wastes (Kang et al. 2008).
In most of the above studies, water quality improved
and the ecological impact was minimized. Moreover, if
the quality of the wastewater is improved enough, it could
be re-used in closed or recirculating systems, for example,
when water is scarce or there is no adequate place to
discharge it (Ruenglertpanyakul et al. 2004) (Fig. 2).
In addition, polyculture allow farmers to produce other
species with commercial value and to increase the eco-
nomical profitability of their farms with little or no finan-
cial investment because most of the costs have already
been met (Purcell et al. 2006). Furthermore, in commer-
cial terms, it has been reported that seafood consumers
will potentially pay an extra 20–30% for products from
polyculture systems (i.e. environmentally friendly foods)
compared with monoculture-farmed products (Ferguson
et al. 2005). For example, oysters and fish produced by
sequential polyculture practices and marketed as ‘special’
oysters and ‘poissons des marais Charentais’, and
achieved a higher commercial value than expected (Bun-
ting 2005). Primavera (1997) suggested that environmen-
tally friendly aquaculture, such as the traditional extensive
polyculture ponds in Indonesia, was one strategy to
improve the cost–benefit ratio of shrimp culture.
There are other species of aquatic organisms with an
acceptable commercial value that are potentially good
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candidates for polyculture. Sea cucumbers, for example,
have high market value (Hamel et al. 2001), but in some
cases their fisheries are restricted because of overfishing
problems (Battaglene 1999); however, some species (Apos-
tichopus japonicas and Isostichopus fuscus) have demon-
strated a high aquacultural potential (Chang et al. 2004;
Mercier et al. 2004). Earthen shrimp ponds provide large
areas suitable for sea cucumbers to thrive and grow ade-
quately, and to be further commercialized as food or to
be restocked into the wild (Pitt & Duy 2004). No special
diet is required as sea cucumbers eat organic and non-
living detrital matter from the sediments (Moriarty 1982).
Likewise, tilapia represents a strong candidate for poly-
culture with shrimp because of its high resistance to
adverse conditions, commercial demand and feeding
habits. Akiyama and Anggawati (1998, 1999) worked in
a ‘direct polyculture’ of shrimp–tilapia and found an
increase in the yield of shrimp and thus an extra eco-
nomic income. Garcia-Perez et al. (2000) reported that
prawn–tilapia polyculture increased economic returns by
21% when cultured in earthen ponds (stocking seven
prawns ⁄ one tilapia m)2; i.e. 1.1 and 7.4 g, respectively).
Some algae that can be used in shrimp polyculture
have an acceptable economic potential. The macroalgae
Kappaphycus alvarezii, which has been co-cultured with
shrimp, has a commercial value because it is the prime
material used to produce carrageenan (Lombardi et al.
2006), a product widely used in the food industry as a
thickening and stabilizing agent. Gracilaria salicornia has
commercial value as an agar source and duckweed (Lemn-
aceae sp.) is used as a high-protein fish feed that thrives
in nutrient-rich waters, such as aquaculture effluent (Rue-
nglertpanyakul et al. 2004). The algae Ulva clathrata,
which has been cultivated in shrimp aquaculture waste
water, shows a high biofiltering efficiency, uptake and
assimilation of nitrogenous compounds (Da Silva-Coper-
tino et al. 2009); in addition, this alga is commonly used
in Asiatic countries as food for human consumption
(Zemke-White & Ohno 1999).
Several reports have indicated that shrimp performance
is enhanced by the presence of a secondary species. In
2006, Martı́nez-Córdova and Martı́nez-Porchas polycul-
tured white shrimp with Crassostrea gigas and Chione
fructifraga and concluded that ‘the presence of mollusks
in the ponds did not have an adverse but a beneficial
effect on the productive performance of shrimp’ (p. 325)
and that C. fructifraga was a good candidate to be
produced in shrimp ponds. Tilapia has been shown to
feed on excess organic matter, improving water quality
and thus increasing shrimp production (Akiyama &
Anggawati 1998). Similar conclusions were drawn by
Wang et al. (1998) in an experimental polyculture of
Chinese shrimp–hybrid tilapia.
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Shrimp polyculture, a sustainable practice
Another benefit of polyculture is its contribution to
enhancing shrimp resistance and ⁄ or protection against
diseases. Many fish have antibacterial, antifungal and
cytotoxic activity in their mucus (Magariños et al. 1995;
Hellio et al. 2002). Antibacterial activity against gram-
negative bacteria was found in aqueous extracts from
the skin mucus of rabbit fish (Siganus fuscescens and
Siganus guttatus). Tendencia et al. (2006) observed that
genetically improved farmed tilapia (GIFT tilapia; 158 g
at 500 g m)3) decreased luminous bacterial counts and as
a consequence increased shrimp (P. monodon) survival.
The same author also demonstrated that other fish, such
as tilapia hornorum (Oreochromis urolepis hornorum)
and milkfish inhibited the growth of luminous bacteria
(Tendencia et al. 2003, 2006). This beneficial effect was
achieved because the mucus of the fish had direct contact
with the water, where the pathogenic bacteria thrive.
Kwon-Jang et al. (2007) found that polyculture of L. van-
namei with river puffer (Takifugu obscures) increased sur-
vival of the shrimp in the presence of white spot syndrome
virus (WSSV) because they ate the moribund shrimps that
were infected by the virus, thus reducing the possibility of
the virus spreading to other shrimps; however, these
researchers reported that there was not the same beneficial
effect when they attempted to use this strategy with
Chinese shrimp (P. chinensis) because Chinese shrimp are
more susceptible. Thus, the addition of some fish to
shrimp ponds may, in some cases, eliminate luminous and
other bacteria and enhance the health status of the shrimp.
Similar results can be achieved with algae. A number of
aquatic plants have chitinase enzymes with antifungal
activity (Shirota et al. 2008). For instance, the seaweed
Gracilaria sp., which has been used in integrated polycul-
ture with shrimp, possesses antimicrobial activity (Sasidh-
aran et al. 2009). Kappaphycus sp., another species used
in shrimp polyculture, has been shown to possess anti-
viral and antibacterial properties (Trono 1999). The sole
presence of Gracilaria textorii in aquatic environments has
been shown to decrease the number of Vibrio sp. (Pang
et al. 2006). However, there is not enough information
about the effect of seaweed on bacterial counts and on
the incidence of viruses in shrimp culture.
Despite the benefits that have been reported with
regard to polyculture practices, this strategy is far from
common among aquaculturists. In Mexico, most farms
are located in the north-west of the country and none of
them carry out polyculture practices; instead they dis-
charge all of their wastewater to the ocean or to estuaries
without previous treatment, and this has caused environ-
mental contamination of the receiving ecosystems (Martı́-
nez-Córdova et al. 2009). There are no reports of penaeid
shrimp farms in Latin America practicing polyculture
strategies on a large scale.
77
M. Martı́nez-Porchas et al.
Disadvantages
Plenty of benefits have been achieved in experimental
shrimp polyculture. However, this activity faces some
problems and disadvantages that have to be considered to
avoid any disaster.
It is important to consider that any beneficial results
can be modified by certain factors, such as the species
involved, the size of the organisms, the density of stock-
ing, stressful conditions, and food quantity and quality
(Pitt et al. 2004; Wang 2007; Da Silva-Copertino et al.
2009). It has been documented that high densities of sub-
ordinate species and stressful conditions diminish shrimp
growth (Muangkeow et al. 2007; Wei et al. 2008).
One of the main problems of polyculture is antagonism
between the species; for example, carnivorous fish can
consume shrimps or shrimps can attack other organisms.
However, according to Chien and Tsai (1985), antago-
nism between the species can be solved by using appro-
priate polyculture, simple engineering (e.g. biomass,
space, oxygen) and proper sequencing of the ponds.
Shrimp polyculture cannot be practiced in farms that
are stocked at intensive densities of shrimp because that
would increase the oxygen consumer biomass (Martı́nez-
Córdova 1999). The use of or the intensification of aera-
tion can solve the problem; however, it is necessary to
evaluate the feasibility and sustainability of this practice.
From an economic viewpoint, polyculture can require
an initial investment, in terms of infrastructure, aeration,
feed and human effort. However, this investment depends
on the type of polyculture, the intensification and the
design of the system (Chien & Tsai 1985; Gonzales-Corre
1988).
Finally, adding a subordinate species risks introducing
pathogens into the culture system. Naylor et al. (2001)
highlighted that a common problem in the aquaculture
industry is the transport and transmission of pathogens
as a result of the use of exotic species.
Despite the disadvantages that have been identified in
shrimp polyculture, it appears that the benefits outweigh
the disadvantages, and in some cases the disadvantages
can be easily solved.
Species requirements
When considering shrimp polyculture it is important to
ensure that the conditions in the shrimp tanks or ponds
meet the biological requirements of the secondary species
that is being co-cultured, otherwise there is a serious risk
of losing all or part of the economic and human effort
invested. For instance, sandfish require a higher water
exchange than the daily 10% carried out in shrimp aqua-
culture and they also require large areas and a place to
78
bury (Pitt et al. 2004). Growth of the mussel Mytilus edu-
lis is severely reduced in polluted environments (Wid-
dows et al. 1981, 1984), which often occurs in shrimp
ponds or ponds effluent. Native species would be good
candidates to filter fed and grow within shrimp ponds or
pond effluent.
It is possible, however, to modify the culture condi-
tions and achieve a range tolerated by two or more
species that are being simultaneously farmed. For exam-
ple, L. vannamei has been shown to thrive well in low
salinity waters with previous and adequate acclimation
(Saoud et al. 2003; Davis et al. 2004). Under these condi-
tions, it is possible to ‘polyculture’ white shrimp with
some fish as reported by Muangkeow et al. (2007) who
cultured L. vannamei with tilapia (Oreochromis niloticus
L.) at a salinity of 24 p.p.t.; in this case adult shrimp nor-
mally live in a salinity of approximately 34 p.p.t. (Martı́-
nez-Córdova et al. 1999) and Nile tilapia is a freshwater
fish (Popma & Masser 1999). The same protocol may be
used for other parameters, such as water temperature,
ionic composition, pH and alkalinity. Ali et al. (2009)
carried out a polyculture with a variety of euryhaline spe-
cies (including fish and crustaceans) and recorded high
survival and production in most of the species cultivated.
Nevertheless, if the environmental conditions of the
culture are modified, the nutritional requirements of the
organisms may also change because growth is dependant
on water temperature, salinity, food quantity and quality
(Dall et al. 1990). For instance, Davis and Saoud (2004)
reported that nutritional considerations have to be taken
into account when rearing shrimp in inland waters with
low salinity.
Another consideration that has to be taken into
account is the feeding habits of the species. It is not
recommended to add species that may be attacked or
devoured by the shrimps or vice versa. Ling (1969) sug-
gested polyculture of the freshwater prawn Macrobrachi-
um rosembergii with non-carnivorous fishes, such as carps
or tilapias, to avoid this problem. When Douglass et al.
(2008) polycultured the grass shrimp Palaemonetes vulga-
ris with the mud crab Panopeus herbstii the survival of
shrimp was completely affected (0%), whereas the sur-
vival of the crab was 100%; however, in a separate mono-
culture, they observed 100% survival of shrimp and 83%
survival of crab. This problem only happens in direct
polyculture and the use of sequential polyculture could
provide a solution.
It is desirable that the supplied feed be consumed by
both species or consumed by the main species, with the
other(s) species consuming the secondary products (feces,
detritus, phytoplankton, zooplankton and metabolites).
Sea cucumbers, for example, feed on nutrient-rich detri-
tus in the ponds of blue shrimp (L. stylirostris) culture,
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not affecting the production parameters of shrimp, but
rather improving the profit as a result of the value of the
sea cucumber (Hammond 1983; Purcell et al. 2006). Jones
et al. (2001) reported that the presence of the Sydney
rock oyster (Saccostrea comercialis) diminished the con-
centration of suspended particles, phytoplankton and bac-
teria in the shrimp effluent, which meant that the waste
nutrients cover, either totally or partially, the nutritional
requirements of the secondary species. Uddin et al.
(2007) documented that prawn feed on the feces of some
fish and calculated a net profit margin of 69% when the
organisms were stocked at a density of 30 000 ha)1 at a
rate of 3:1 (tilapia:prawn).
Lombardi et al. (2006), experimented with cage poly-
culture of Pacific white shrimp and Philippines seaweed
(Kappaphycus alvarezii) and obtained production parame-
ters as high as 3.23 and 23.70 kg m)2 year)1 for shrimp
and seaweed, respectively; in this trial shrimps were
stocked inside the cages to avoid grazing on the seaweed.
Jones et al. (2001) observed that the marcoalgae Gracilar-
ia edulis absorbed the dissolved nutrients produced in
shrimp culture tanks.
It is also important to consider that the subordinate
species may be a vector for disease. Some of these species
can be exotic and can bring pathogen viruses and para-
sites that may affect the shrimp, while the exotic species
is immune (Sahul-Hameed et al. 2002). Under that sce-
nario, the yield of the shrimps and the economic profit
would decrease instead of increase; for that reason, it is
recommended that the subordinate species be analyzed
for viruses, bacteria and parasites before they are added
into the shrimp culture.
The size of the species at the time of stocking is an
important aspect, particularly in direct polyculture. Pitt
et al. (2004) reported that shrimp–sandfish polyculture was
not profitable because big shrimp (>1 g) caused the death
of small sandfish ( 1 g). Better results were achieved in
polycultures of ‘big shrimp–big sandfish’, ‘small shrimp–
big sandfish’ and ‘small shrimp–small sandfish’. In the case
of bivalves size is apparently not a factor that can affect
shrimps thriving; however, in organisms such as fish, ‘size’
and ‘species’ may affect shrimp performance.
Stocking rate is another concern in polyculture
management. Tian et al. (2001) worked with ‘Chinese
shrimp–red tilapia–constricted tagelus’ polyculture (2 cm
shrimp, 150 g tilapia and 3 cm tagelus) and found the
best stocking rate to be 7.2 shrimp m)2, 0.08 tilapia m)2
and 14 tagelus m)2, whereas other combinations (0:20,
0.12:10, 0.16:7 and 0.24:0 tilapia:tagelus m)2) showed a
lesser yield. Wang et al. (1998) reported an optimum
stocking density of six Chinese shrimp m)2 and 0.32 red
tilapia m)2. Apparently shrimp polyculture systems per-
form well at mid densities because overstocking compli-
Reviews in Aquaculture (2010) 2, 73–85
ª 2010 Blackwell Publishing Asia Pty Ltd
Shrimp polyculture, a sustainable practice
cates the management and is not sustainable in the long
run (Yi & Fitzsimmons 2004), although further research
in this area is needed.
System design
The design of shrimp polyculture units will have an
impact not only on the production results, but also on
the ecology of the involved ecosystems because ecological
interactions between two or more species may have a syn-
ergic or antagonistic effect. It is difficult to establish spe-
cific criteria for designing a successful polyculture system
because of the high number of possibilities and the pleth-
ora of papers reporting successful results using diverse
systems with different engineering. However, in general
terms, it is crucial that the systems provide the environ-
mental requirements of the cultured organisms and avoid
where possible any type of negative interaction between
the species. There is not a general or universal design for
shrimp polyculture, rather the designs may vary according
to the compatibility and requirements of the particular
species (Chien & Tsai 1985; Troell et al. 2009). In this
section we outline the most important aspects that need
to be considered.
Monotrophic and multitrophic polyculture
Multitrophic polyculture is a better option compared
with monotrophic polyculture because in monoculture
the organisms share the same biological and chemical
processes, which in turn may lead to a decrease in envi-
ronmental quality. In contrast, multitrophic polyculture
combines the feeding habits of different organisms, main-
taining an ecological equilibrium within the system (Cho-
pin 2006; Shimoda et al. 2006). As stated, this practice
promotes the formation of nutrient cycling, impeding the
accumulation of wastes or toxic components. For exam-
ple, in a shrimp–macroalgae–bivalve polyculture, shrimp
feed on an artificial diet, phytoplankton, zooplankton and
other sources, whereas the seaweed absorb the nitroge-
nous wastes produced by the shrimps and the bivalves
feed on the organic particulate matter produced by the
feces and the non-consumed pellets.
Food administration and accessibility
To achieve success in a polyculture venture one of the
first concerns of the system designer is to ensure accessi-
bility to food for both (or more) species being farmed.
Competition for food has been a problem in shrimp poly-
culture systems when free-swimming fish are present
(Dos Santos & Valenti 2002) because these organisms are
faster swimmers than shrimps and monopolize the feed,
79
M. Martı́nez-Porchas et al.
and shrimp growth is negatively affected. Muangkeow
et al. (2007, p. 365) affirmed that ‘mixed shrimp–tilapia
polyculture systems without partitioning have not suc-
ceeded due to the decreasing growth and yield of shrimp’;
therefore, it is suggested that the fish or the shrimps are
maintained inside cages to ensure food accessibility. In a
similar experiment, Martı́nez-Córdova and Peña-Messina
(2005) carried out a direct ‘white shrimp–blue shrimp’
monotrophic polyculture, in which the growth of white
shrimp was optimized, but the growth and survival of
blue shrimp were poor, probably because of its higher
susceptibility to adverse conditions and because the white
shrimp monopolized the food (observed by stomach con-
tents). Tian et al. (2001) experimented with the poly-
culture of Chinese shrimp with constricted tagelus
(Sinonovacula constricta Lamarck) and hybrid tilapia
(Oreochromis mossambicus · O. niloticus) confined in
cages and showed that the tilapia retained approximately
3% of the total input of nitrogen and that growth of the
shrimps was acceptable. An alternate and suggested solu-
tion is sequential polyculture or integrated systems (San-
difer & Hopkins 1996) where subordinate species thrive
in separate units supplied with shrimp effluent. Erler
et al. (2004) demonstrated that tilapia and grey mullet
can be cultivated in shrimp pond effluent and, moreover,
that denitrification increased, removing up to 21% of the
total nitrogen. In addition, some reports have exposed
the theory that the wastewater of some cultured fish con-
tains valuable nutrients, such as organic and suspended
matter (Jiménez-Montealegre et al. 2002; Al-Hafedh et al.
2003); this was further demonstrated by Kuhn et al.
(2008) when they fed shrimps (L. vannamei) with a com-
mercial diet complemented with ‘bio-flocs’ generated
from tilapia effluent and obtained better production
parameters compared with shrimps fed only on a com-
mercial diet. They concluded that this strategy offered an
alternative and sustainable approach to integrated shrimp
farming. Other authors that have worked with partitioned
systems have demonstrated their efficacy as wastewater
bioremediators and yield improvers (De-shang & Shuang-
lin 2000; Guocai et al. 2000a).
Partitioning and direct polyculture
According to efficacy results, partitioning polyculture sys-
tems (cage-cum-pond or sequential polyculture) appear
to be a better option than direct polyculture; these sys-
tems also have the advantage of saving time at harvest
and biometry’s activity because it is not necessary to sepa-
rate one species from the other or to disturb the different
species that are being cultured.
However, if direct polyculture is the only option, it is
necessary to consider that some penaeid shrimps exhibit
80
cannibalistic behavior and may attack other species
(Martı́nez-Córdova et al. 1999); therefore, it is necessary
to provide hideout places for secondary species. In this
context, it has been reported that the presence of shrimp
affected the survival and growth of the sandfish Holothu-
ria scabra (Bell et al. 2007) probably because of the lack
of sand ⁄ sediment for burying (Mercier et al. 1999, 2000).
In contrast, Purcell et al. (2006) recorded high survival of
shrimp (79–90%) and sandfish (73–100%) in an experi-
mental polyculture where sandfish buried into the sedi-
ments in the early morning and thus obtained protection
from harassment by the shrimp. Pitt et al. (2004)
reported successful results in the production parameters
of blue shrimp and sandfish and concluded that the sur-
vival and growth of sandfish were better in the presence
of shrimps, perhaps because the shrimp improved the
benthic conditions and the sandfish had the opportunity
to bury.
Increased area
Polyculture implies an increase in the density of organ-
isms in the system, all of which use the same space, oxy-
gen and feed. This condition may eventually become a
serious problem and make it necessary to increase the
farming area. An alternative to increasing the area with-
out augmenting the farm size is the addition of substrates
into the pond.
The use of artificial substrates, either artisanal or
commercial (e.g. Aquamats), has shown good results in
crustacean–fish polyculture because the contact surface
is increased, promoting biotic communities such as
phytoplankton, zooplankton and periphyton, which
contribute to the nutrition of the shrimp and the co-
cultured species. This practice also reduces turbulence
and water turbidity, improving the water quality. In a
prawn–tilapia polyculture, Uddin et al. (2007) placed
vertical bamboo sticks within the experimental ponds
and promoted periphyton growth, which served as a
food source for both species. Erler et al. (2004) reported
an increase in solids settlement and water quality in
shrimp effluent when fish and bivalves were cultivated
and aquamats were installed; nitrogen removal increased
threefold (2.4%) compared with treatments without
aquamats (0.8%).
Oxygen management
The addition of a subordinate species into the shrimp
culture system increases the oxygen demand because the
total biomass of the system is increased; in such a case
emergency or supplemental aeration is required (Danaher
et al. 2007). Guocai et al. (2000b) argued that phyto-
Reviews in Aquaculture (2010) 2, 73–85
ª 2010 Blackwell Publishing Asia Pty Ltd

plankton and sediment demand high amounts of oxygen
in shrimp polyculture systems. Nonetheless, subordinate
species can consume organic matter and microalgae,
reducing the oxygen demand of the entire system (Tian
et al. 2001). Regarding seaweed as a candidate for sub-
ordinate species, it is necessary to consider that high densi-
ties of seaweed can produce hypoxia in the early morning
(Lanza-Espino 1998). Algae as subordinate species can be
used in partitioning polyculture, with preference to
‘sequential polyculture’, and the plants should be placed
at the end of the system (i.e. in the effluent-receiving
ponds). In addition, the presence of organisms within the
ponds may affect the growth of the plants because the
turbidity of the water is increased owing to the distur-
bance of sediment by either shrimp or fish movements;
Phang et al. (1996) stated that turbidity had a negative
effect on the growth of seaweed.
Aeration or oxygenation units, either for normal
(depending on the biomass) or emergency cases, should
be included in all system designs.
Increase in natural productivity
Another consideration in polyculture systems is improve-
ment of the environmental and water-quality conditions
within the pond to enhance the productivity of natural
food and therefore increase yield production and trans-
form excess nitrogen into non-antagonistic (organic
matter and toxic chemicals) subproducts for shrimp
(phytoplankton and zooplankton). For example, Marti-
nez-Cordova and Enriquez-Ocaña (2007) introduced
enclosures (without shrimp) into shrimp farm pond and
effluents and found that the total benthic fauna, particu-
larly polychaeta m)2, increased either in the pond or dis-
charge lagoon, and had a positive impact on shrimp
growth performance; thus, enclosures provided natural
feed for shrimps and reduced the consumption of artifi-
cial feed, which in polyculture perspectives would benefit
subordinate species such as fish and reduce feeding costs.
Several studies (Tacon et al. 2002; Cuzon et al. 2004;
McLean et al. 2006) have demonstrated that shrimp per-
formance, in terms of health, growth and efficiency in
nitrogen utilization, is enhanced when microbial flocs,
algae and other natural biota are generated within the
culture system.
Engineering
With regard to the engineering design, it may vary
according to, for example, the species requirements, the
settling environment and economic resources. As general
characteristics of concern, oxygen administration, water
flow and pond logistics (sequence, places to install the
Reviews in Aquaculture (2010) 2, 73–85
ª 2010 Blackwell Publishing Asia Pty Ltd
Shrimp polyculture, a sustainable practice
cages) need to be taken into consideration. However,
every design involves particular engineering.
There are plenty of diverse system designs; however,
despite all of the design characteristics being taken into
account, successful or unsuccessful polyculture results do
not depend only on technical aspects, but also on the
social and economic factors that prevail in the place in
which the polyculture is (or planned to be) practiced
(Stewart & Seijo 1994; Ponce-Marbán et al. 2006).
Conclusions
The shrimp aquaculture industry faces several problems,
including contamination caused by farm effluent. Polycul-
ture appears to be an adequate alternative to minimize or
eventually solve these problems. However, polyculture is
not commonly used in the shrimp industry. There are
three general types of polyculture: direct, cage-cum-pond
and integrated polyculture. Each one with two variants:
monotrophic and multitrophic.
Shrimp polyculture is a promising industry and has been
shown experimentally to improve yield, decrease the envi-
ronmental impact, re-utilize wastes and reduce the inci-
dence of pathogen organisms in shrimp aquaculture. In
addition, different by-products with a commercial value can
be obtained from the subordinate species. However, there
are problems and considerations that need to be solved
before practicing polyculture, such as economic and human
effort investment, compatibility of the species (feeding hab-
its and environmental requirements), food competition,
space, size of the organisms and decreasing oxygen levels.
Many species from different trophic levels, such as fish,
mollusks and plants, show aquaculture potential for use
as subordinate organisms in shrimp farming. However, it
is important that these species fit with the environmental
requirements of the shrimp.
Partitioning and multitrophic polyculture appears to be
the best option for this practice because possible antago-
nistic activity among the species is diminished when they
are separated by different units or barriers (cages).
The design of the culture system should be cost effec-
tive and provide an optimal environment for the species
that are being cultured to thrive. However, further
research is needed in this field that examines new species,
densities and systems. It is also necessary to make aware
and to encourage farmers to try this activity to reduce the
environmental impact resulting from effluent discharge.
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