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Reviews in Fisheries Science & Aquaculture

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Integrated Freshwater Prawn Farming: State-of-the-Art and Future Potential

Helcio L. A. Marques, Michael B. New, Marcello Villar Boock, Helenice PereiraBarros, Margarete Mallasen & Wagner C. Valenti

To cite this article: Helcio L. A. Marques, Michael B. New, Marcello Villar Boock, HelenicePereira Barros, Margarete Mallasen & Wagner C. Valenti (2016) Integrated Freshwater PrawnFarming: State-of-the-Art and Future Potential, Reviews in Fisheries Science & Aquaculture,24:3, 264-293, DOI: 10.1080/23308249.2016.1169245

To link to this article: http://dx.doi.org/10.1080/23308249.2016.1169245

Published online: 20 Apr 2016.

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Integrated Freshwater Prawn Farming: State-of-the-Art and Future Potential

Helcio L. A. Marquesa, Michael B. Newb, Marcello Villar Boocka, Helenice Pereira Barrosc, Margarete Mallasenc,y, andWagner C. Valentid

aAquaculture Center, Fisheries Institute, Sao Paulo State Secretariat of Agriculture and Food Supply, S~ao Paulo, Brazil; bFreshwater PrawnFarming Research Group, CNPq, Brazil, Marlow, Bucks, UK; cCenter for Continental Fish, Fisheries Institute, Sao Paulo State Secretariat ofAgriculture and Food Supply, S~ao Paulo, Brazil; dBiosciences Institute and CAUNESP, CNPq., S~ao Paulo State University UNESP, S~ao Paulo, Brazil

ABSTRACTIntegrated aquaculture can be defined as aquaculture systems sharing resources with otheractivities, commonly agricultural, agroindustrial, and infrastructural. Freshwater prawns are excellentoptions for integration, since they are omnivores and can therefore take advantage of a wide rangeof feed residuals, either from aquatic or terrestrial species. Furthermore, due to their benthic habit,they have a well-defined spatial distribution in the environment, thus favoring interaction withvarious species of fish, other animals, and even with plants. The integrated farming of freshwaterprawns includes different culture systems, such as polyculture and coculture with other aquaticspecies, rice-prawn culture, hydroponics, and integration with terrestrial animals and plants. Ourreview includes a worldwide perspective on the main commercial integrated systems involvingfreshwater prawns, the present status of research on integrated freshwater prawn production andthe main opportunities for integrated freshwater prawn farming in a world that is moving towardsustainability. The review continues by providing a brief summary of the future prospects for thisform of aquaculture. Finally, we conclude that integrating freshwater prawn farming with otheraquaculture and farming activities has considerable potential as a means of increasing foodproduction in a sustainable fashion.

KEYWORDSfreshwater prawns;Macrobrachium; polyculture;coculture; integration;potential; IMTA

Introduction

The term freshwater prawn, sometimes referred to asfreshwater shrimp is a general designation for carideancrustaceans, which spend at least the juvenile and adultphases in freshwater. Most species farmed around theworld or used in aquaculture experiments belong to thegenus Macrobrachium. This genus is tropical and nativeto Asia, Africa, and America (Holthuis and Ng, 2010).Until recently, most commercial culture had been basedon the giant river prawn Macrobrachium rosenbergii.Nevertheless, >250,000 t of the oriental river prawnMacrobrachium nipponense were produced in China in2013, the culture of the monsoon river prawn Macro-brachium malcolmsonii has started in India and Pakistan,and the monkey river prawn Macrobrachium lar inVanuatu (FAO, 2015). Many other species are underresearch for aquaculture purposes, including the Ama-zon river prawn Macrobrachium amazonicum and thepainted river prawn Macrobrachium carcinus in theAmericas, and the African river prawn Macrobrachium

vollenhovenii (see Kutty and Valenti, 2010 for a review).Recently, freshwater prawn farming has become a majorcontributor to global aquaculture, both in quantity andvalue (New, 2010). In 2013, 571,152 t of freshwaterprawns were produced globally, which makes freshwaterprawn farming an industry valued at the farm-gate atUS$ 3 billion/year (FAO, 2015).

The term “integrated aquaculture” has been fre-quently misunderstood. In scientific papers and in thedocuments of government agencies or private enter-prises, there is often a confusion between the terms “inte-grated aquaculture,” “sustainable aquaculture,” “organicaquaculture,” “responsible aquaculture,” and “sociallyand/or environmentally responsible aquaculture.” Theseexpressions are quite distinct, although they have someelements in common. For example, a fish monoculturemay be sustainable, or organic, or environmentallyresponsible, despite not being an integrated aquaculture.On the other hand, an integrated aquaculture systemdoes not have to be organic and sometimes, it may not

CONTACT Wagner C. Valenti valenti@clp.unesp.br Universidade Estadual Paulista, Instituto de Biociencias, Caixa Postal 73601, 11380-972, S~ao Vicente,SP, Brazil.

Color versions of one or more of the figures in this article can be found online at www.tandfonline.com/brfs.yIn memoriam of Margarete Mallasen.© 2016 Taylor & Francis

REVIEWS IN FISHERIES SCIENCE & AQUACULTURE2016, VOL. 24, NO. 3, 264–293http://dx.doi.org/10.1080/23308249.2016.1169245

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be sustainable or may not even be conducted in anenvironmentally and socially friendly way.

According to the Food and Agriculture Organization(FAO, 2013), integrated aquaculture is defined as “aqua-culture systems sharing resources (e.g., water, feeds,management, etc.) with other activities, commonly agri-cultural, agroindustrial, and infrastructural (wastewater,power stations, etc.).” Zimmermann et al. (2010) defineintegrated aquaculture as “systems that combine aqua-culture with the farming of terrestrial animals andplants.” These authors distinguish it from the term “pol-yculture,” used to name the art of growing two or morecompatible aquatic species together in a single pond.These definitions have been largely used in aquaculture.In recent years, the expression “integrated multi-trophicaquaculture” (IMTA) has become popular to describethe farming of species from different trophic levels andwith complementary ecosystem functions. In IMTA,uneaten allochthonous feed, wastes, nutrients, and by-products from one species can be caught by other speciesand incorporated in biomass (Chopin et al., 2012; Cho-pin, 2013). This concept also includes polyculture. Thus,in our review, we consider fish-prawn polyculture as aform of integrated system, since it shares the availableresources in a complementary way, unlike some polycul-ture systems, which involve two or more fish species thatcompete for the same resources, rather than share themcomplementarily.

Sharing natural or allochthonous resources is a way ofincreasing their usage efficiency. Thus, integrated aqua-culture systems are supposed to optimize the use ofresources, contributing to the conservation of finite natu-ral assets. In addition, it is possible to exploit the syner-gistic interactions between the farmed species, leading tobiomitigation processes. Consequently, integrated aqua-culture generally shows higher sustainability than othersystems, as will be shown in this review.

Integrated freshwater prawn farming includes variousfarming systems, such as fish-prawn and rice-prawn cul-ture, aquaponics, and integration with terrestrial animalsand plant crops. All these forms of integration can beprofitable. The decision about implementing an inte-grated system and the choice of integration type dependon the characteristics of the farm, the technical and eco-nomic viability of the integration and the existence oftechnology and market niches for all the species pro-duced. A previous review of integrated freshwater prawnfarming was providing by Zimmermann et al. (2010). Inthis paper, available information has been updated, andnomenclature has been standardized. The current reviewfocuses mainly on the great advantages of introducingfreshwater prawns into aquafarms previously operatedusing plants or other animal species. For the purposes of

this review, polyculture is defined as the rearing of twoor more species (e.g., prawns and tilapia) in water bodieswithout physical separation, whereas coculture is therearing of two or more species in the same water bodybut incorporating physical separation (e.g., one speciesin cages).

Benefits of using freshwater prawns inintegrated aquaculture

Freshwater prawns are a good option for integrated sys-tems, since they are omnivores and detritivorous andhave a benthic habit. Thus, they can take advantage of awide range of feed wastes, either from aquatic or terres-trial species, which fall through the water column bygravity and sediment at the bottom of the rearing sys-tems. Furthermore, they have a well-defined spatial dis-tribution in the environment, and occupy a slender layerin the bottom of the tridimensional space of aquatic sys-tems. This avoids competition with various species offish, and even allows association with plants.

The addition of freshwater prawns to a culture systemmay add value to the crop. Generally, fish, rice, and otherfreshwater organisms have low market prices. Con-versely, freshwater prawns attain a very high price; forexample, the global average in 2013 was US$ 5.23/kg(FAO, 2015). The inclusion of a higher value species inan aquaculture system can yield a significantly increasedincome for the farmer, even when an increase in produc-tivity does not occur (Zimmermann et al., 2010). Thus,freshwater prawns provide high income even whenreared at low densities in fish ponds or rice fields,increasing total crop value. In addition, their use in inte-grated systems requires minimum alterations in facilitiesand management. New (2002) stated that the introduc-tion of prawns in paddy fields does not reduce the pro-duction of rice; for example the profits from Vietnameseintegrated rice-prawn culture can be two or three timesgreater than from rice monoculture.

Generally, the performance of freshwater prawns isaffected only by their own stocking density and manage-ment. It has been observed that the density of the fishstocked together and the feeding or fertilizing regimesused for the costocking species has only a minor effecton freshwater prawns (Wohlfarth et al., 1985; Ara�ujoand Portz, 1997). Thus, rearing prawns in low densitywith fish or in rice paddies generally results in the pro-duction of large-sized prawns with high market value.

The introduction of freshwater prawns into other cul-ture systems may even improve the performance of thecostocking organism. Uddin et al. (2006) observed a pos-itive effect of polyculture on the growth of both fish andprawns. In addition, rice production in Bangladesh has

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been observed to increase in an integrated system(prawn-fish-rice) because the prawns and fish improvesoil fertility and promote an integrated pest management(Ahmed et al., 2010; 2014).

In conclusion, there is great potential for rearingfreshwater prawns using the bottom of uncountableareas of tropical and subtropical fish ponds and ricepaddy fields around the world. This provides an oppor-tunity for fish and rice farmers to increase productionand profit with minor extra investment. Furthermore,this increased food production and profit is attainedwithout any additional land use, saving the environment.Therefore, the use of freshwater prawns in integratedsystems has a great potential to increase food security,improve local and global economy, and increase thesocial, economic and environmental sustainability ofaquaculture systems. In the following sections of thisreview, the various systems in which freshwater prawnsmay be incorporated are described, and the current sta-tus of practical integration and research is considered.

Production systems suitable for freshwaterprawn integration

Fish-prawn polyculture

New (2002) listed some synergistic beneficial effectsresulting from the inclusion of freshwater prawns in pol-yculture systems. These include the maintenance ofmore stable dissolved oxygen levels, reduction of preda-tors, increased feeding efficiency, increased pondproductivity, and a potentially increased total crop valuethrough the inclusion of a high-value species. Zimmer-mann et al. (2010) highlighted the more rational use ofthe pond because of the different feeding habits of thespecies and the additional income from freshwaterprawns, which have high market value. The sameauthors, however, cautioned that proper knowledge ofthe feeding habits of the species that are cultured withthe freshwater prawns is essential. As prawns are tropicalanimals, the fish to be reared with them must be adaptedto warm waters; thus, salmonids and other temperatespecies are excluded.

The commonest form of integrated fish-prawn cultureis within the same pond, where both prawns and fishgrow together freely. The fish used should be pelagic,non-carnivorous and non-aggressive. In addition, theculture cycle duration of the fish should be similar toprawns, to allow the stocking and harvesting of all spe-cies simultaneously. The freshwater prawns walk/swimand eat on the bottom, whereas pelagic fish mainly swimin the water column, feeding on floating artificial diets oron plankton, suspended detritus, or aquatic plants. Thus,

available space and food can be optimized. Some fishspecies, such as Nile tilapia (Oreochromis niloticus), silvercarp (Hypophthalmichthys molitrix), and bighead carp(Aristichthys nobilis) are filter feeders or actively eatplankton, whereas common carp (Cyprinus carpio) areomnivorous and grass carp (Ctenopharyngodon idella)are primarily herbivorous. Thus, these species in polycul-ture with prawns take advantage of all these feedingniches (Malecha et al., 1981). In addition to increasingtotal production in the pond, the fish in polyculture pre-vent the excessive growth of phytoplankton, zooplank-ton, macrovegetation, and filamentous algae (Cohenet al., 1983).

In polyculture, prawns can eat tilapia feces andunused fish feed (Santos and Valenti, 2002), while tilapiafilter phytoplankton (Perschbacher and Lorio, 1993) andzooplankton (Ibrahim et al., 2015), reducing the risk oflow dissolved oxygen levels at night. In addition, prawnbioturbation at the pond bottom returns nutrients to thewater column (Kimpara et al., 2011), enhancing phyto-plankton production and consequently the natural feedavailable for the tilapia. Prawn larvae hatched insideponds may also be filtered within the plankton and eatenby tilapia (Guerrero and Guerrero, 1979). When kept insmall tanks, tilapia (mainly fry) eat prawn postlarvae,while large prawns may damage tilapia. Such agonisticbehavior can be avoided by coculturing (confining onespecies, e.g., in cages) and is considerably reduced in pol-yculture in large ponds because of the differing spatialdistribution of the species and/or size and stocking den-sity management. Tilapia and prawns exploit differentniches in the pond environment, show positive syner-gism and low antagonistic interactions; thus, they can becultivated together with success.

Generally, prawns are stocked in polyculture systemsat lower densities than in monoculture, whereas the fishare stocked at densities similar to their monoculture.Therefore, prawn productivity is reduced in polyculture,whereas the fish productivity is similar to that frommonoculture. In this case, the emphasis of the polycul-ture is on the production of fish, and thus fish arereferred to as the “main” species in the system (Cohenet al., 1983). Usually, only the fish are fed with commer-cial pellets, while the prawns feed on waste feed, fishfeces, and/or on benthic organisms that grow on thepond bottom, stimulated by nutrients present in the fecesand excreta of the fish (Santos and Valenti, 2002). Theprawns must be stocked first, followed 7–10 days later bythe stocking of fish. This simple procedure greatlyreduces the predation of juvenile prawns by fish (Santosand Valenti, 2002).

A promising form of fish-prawn culture is the cocul-ture of caged fish and free prawns in the same pond

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(Rouse and Stickney, 1982; Tidwell et al., 2000, 2010;Danaher et al., 2007) (Figure 1). The fish receive artificialfeed, but the prawns may or may not be fed, dependingon their stocking density. At low densities (< »80 gm¡2), prawns are normally not fed. In addition, the cagesand pens where the fish are farmed provide shelter andsubstrates for the prawns, as well as additional feed dueto the periphyton that grows on those structures. Har-vesting is greatly facilitated since it is not necessary toseparate the species manually (Danaher et al., 2007), andfish can be harvested regardless of the presence of theprawns. Thus, the range of species that can be used inthis system is not constrained, and can potentiallyinclude carnivorous fish and species with a culture cycleshorter than that of freshwater prawns. Species with along rearing cycle should be avoided because it is difficultto harvest the prawns before the fish. The emphasis ofthis system may be placed either on fish or prawns,depending on the stocking density and management ofeach species adopted by the farmer.

Rearing caged prawns and freely swimming fishwithin ponds is feasible, but not common. Prawn growthinside cages may be reduced after maturation (Marqueset al., 1998). Nevertheless, promising results wereobtained by Cuvin-Aralar et al. (2007) rearing M. rose-nbergii in cages in a lake in the Philippines. In this study,juveniles (0.4 g) were cultured in 2.5 m £ 1 m £ 1 mcages at replicated stocking densities of 15, 30, 60, and90 prawns m¡2 of cage bottom. Mean sizes at harvestafter 5 months of culture ranged from 14.3 g for thehighest stocking density to 26.3 g for the lowest. Meansize at harvest and daily growth rate were significantlyinfluenced by stocking density, with those at the loweststocking density showing significantly better growth.Survival substantially decreased in the highest stocking

density (55.3, 54.0, 52.7, and 36.9% for 15, 30, 60, and 90prawns m¡2, respectively). These authors noted that theproduction from M. rosenbergii cultured in cages wascomparable to or even higher than those reported frompond culture, given that the stocking densities used inthis study were generally higher than in ponds. In addi-tion to grow-out, the freshwater prawn nursery phasecan be successfully carried out in cages inside pondsusing very high postlarval (PL) densities, such as1200 m¡2, with high survival and profit (Marques et al.,2000; 2010; 2012). Cages may decrease the swimmingspace available for pelagic fish, such as tilapia, but theyprovide extra feed for the fish because of the periphytonthat develop on the cage structures and netting. On theother hand, cages may simply occupy vacant space in thepond culture of benthic fish, such as catfish.

Rearing fish at low densities in ponds where freshwa-ter prawns are stocked at the same densities as in mono-culture has also been practiced. In this case, fishmaintain the ecological balance and good water qualityinside the ponds without disturbing the prawns (Cohenet al., 1983). Fish are much more efficient than prawns inseizing feed pellets, and thus increasing their density willaffect prawn feeding, decreasing production (New,2002). The marketable value of fish is normally lowerthan prawns; thus this form of polyculture does notgreatly increase profits, but may be a way of improvingpond water quality, controlling some predators or inva-sive plants, providing an extra income through the saleof fish, or providing extra food for family consumption.

Rice-prawn culture

The harvest of fish and prawns from inundated rice fieldsis an ancient and traditional practice in many Southeast

Figure 1. Coculturing of tilapia in cages placed inside prawn ponds in Brazil (photo by Helcio Marques).

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Asian countries (Das, 2002; Galib, 2010). The first fresh-water prawn cultivation in rice fields appeared in Bangla-desh between the late 1970s and the mid-1980s (Ahmedet al., 2010). During the last two decades, integratedprawn-fish-rice farming has been expanded in Asia,mainly because the export potential of M. rosenbergiiand its high market value (Ahmed et al., 2010; 2014).Wild-caught juvenile prawns have been replaced by juve-niles produced in hatcheries; allochthonous feedingoccurs and water quality control is introduced, makingthe rice-prawn culture a commercial activity rather thanan artisanal practice (Phuong et al., 2006).

There are several interactions between prawns andrice in rice-prawn systems. The paddy fields provideflooded space, shade, and shelter against prawn preda-tors, keeping the water temperature at tolerable levels inthe hot summer months (Ahmed and Garnett, 2010). Inturn, prawn and fish wastes increase the amount oforganic material in the rice fields, decreasing the neces-sity for fertilization. Prawns also contribute to the partialcontrol of some weeds, insects, and pests. If prawns arefed, feed waste decomposition results in the release ofnitrogen, phosphorus, and other elements important forfertilization purposes in rice culture. Rice paddies do notprovide a totally safe environment for prawns andgrowth reduction or high mortality rates may occurbecause of temperature oscillations and predation(Boock et al., 2013, 2016).

According to Phuong et al. (2006), there are two mainforms of rice-prawn farming: rotational culture (alter-nate culture) and simultaneous culture (integrated farm-ing). The choice depends on farmer preference or thesite characteristics. In the rotational system, prawns arereared in the monsoon season and rice in the dry season.The flooded rice fields are stocked with prawn PL orjuveniles at densities of »3–12 prawns m¡2 and harvest-ing occurs after 6–8 months, resulting in large prawnswith 50–110 g body weight. Productivity is about 900 kgha¡1. In the simultaneous culture system, rice fields are

stocked with juvenileM. rosenbergii (2–3 cm) at densitiesfrom 1.5 to 5 m¡2 for 2 to 2.5 months (Figure 2). Simul-taneous culture is performed during the monsoon season(summer-autumn rice crop) and, after that, only rice iscultured in the dry season. Productivity is low, rangingfrom 40 to 500 kg ha¡1, due to the small size of prawnsat harvest and to low survival rates.

In the simultaneous system, the rice paddies must beadapted for prawn stocking. A trench (called a harvestchannel) with a depth of 80–100 cm must be dug sur-rounding the rice field or along the dike near to the wateroutlet point (Hung 2001) (Figures 2 and 3). This ditch isessential for the harvesting of both prawns and ricebecause most prawns shelter in the channel when therice field is drained (Figure 4A and B). After the rice hasbeen harvested, the level of water may be increased againto allow the prawns to eat the leftover by-products of therice culture (Figure 4C). In addition, the level of the dikesmust be increased to enable a raised level of water to»30 cm and individual systems of water inlet and outletneed to be built (Hung, 2001). Screens are provided to

Figure 2. Rice-prawn simultaneous culture. Prawns are stocked in the rice paddies together with the rice plants.

Figure 3. Rice-prawn culture in India showing the rice paddy andthe lateral trench (photo by Helcio Marques).

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prevent the escape of prawns. In Bangladesh, these modi-fied rice fields are referred to as ghers (Ahmed et al.,2010). The gher system has also been described as anenclosure made for prawn cultivation by modifying ricefields through building higher dikes around the field andexcavating a canal inside the periphery to retain waterduring the dry season (Ahmed and Garnett, 2010).

The gher system is a combined form of agriculture andaquaculture, incorporating the joint operation of threeenterprises: prawn, fish, and rice production (Rahmanet al., 2011). In addition, a variety of small-scale vegetablesand fruits are produced on the dikes. This includes carrot,tomato, onion, mustard, long yard bean, spinach, pea,potato, sweet gourd, cucumber, chilli, and some short-cycle fruits such as banana, papaya, and guava (Ahmedet al., 2008c; Barmon, 2014). Gher systems can use eitherfreshwater, when only freshwater prawns are farmed, orbrackishwater when marine shrimp are included (Barmonet al., 2006). This system had been expanding rapidly inthe coastal areas of Bangladesh and has had significantimpacts on the agricultural characteristics and economyof Bangladesh (Rahman et al., 2011).

Aquaponics

Aquaponics links recirculation aquaculture with hydro-ponic vegetable, flower, or herb production (Diver and

Rinehart, 2010). In this system, nutrient-rich effluentfrom aquaculture is used for hydroponic vegetable pro-duction. In turn, plants and rhizobacteria remove ammo-nia, nitrates, nitrites, and phosphorus from the waterbefore it is recirculated back into the prawn tanks. Inaddition, prawns control pests and remove wastes fromthe tanks. Aquaponics may be a great opportunity forurban and suburban family food production because itrequires little space. The main components of aquaponicunits are a hydroponic container, where plants grow, aprawn-fish tank, a mechanical filter and a biofilter.Pumps are used to circulate water throughout the variousunits.

According to Somerville et al. (2014) prawns arevery suitable for inclusion in aquaponics systemssimultaneously with mid-water fish. They consumeuneaten fish feed and suspended and benthic organicwastes. Thus, they accelerate organic matter decom-position and contribute to the health of the system.Some systems with tilapia have been tested withsuccess.

The selection of plant species adapted to aqua-ponics is necessarily related to prawn stocking densityand the consequential nutrient concentration of efflu-ents. Lettuce, herbs, and greens (spinach, chives, basil,and watercress) have low to medium nutritionalrequirements and are well adapted to aquaponic

Figure 4. Rice-prawn culture crop management. Prawns shelter in the refuge (trench) when the rice field is drained (A). Thus, they canbe harvested or selected and rice plants can be managed or cropped (B). After the rice has been cropped, the level of water may beincreased again to allow the prawns to eat the leftover by-products of the rice culture (C).

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systems. Nevertheless, the nutrient composition of theeffluent may not fully match the requirements ofthese plants; thus, an additional nutritive solutionmay be required.

Other crops

The simplest and earliest form of integration with thefarming of terrestrial plants other than rice is the use ofpond banks (bunds) for planting small trees such asbananas, coconut, and papaya. Such integrated systemsoptimize the space available in paddies and pond layoutsand provide an extra income derived from fruit produc-tion. Nevertheless, New (2002) stated that large trees orplants with extensive root systems planted on top of thebunds may damage them and cause leakage. Prawn pondeffluent can be used for the fertilization/irrigation ofcrops such as greens, vegetables or fruits (New, 2002).Effluent water is rich in nutrients, such as nitrogen andphosphorus and can thus increase soil fertility, especiallywhen crops are able to absorb nutrients rapidly, as is thecase with vegetables and herbs.

Livestock

The integration of prawn culture with livestock produc-tion enables the use of animal feces and excreta for pondfertilization and provides other benefits. This type ofintegration may include pig, cattle, goat, sheep, chicken,and duck production. In the case of pigs, their feces maybe used directly as feed, because of they contain 70%digestible food (Tripathi and Sharma, 2001b; Hung,2001). In addition, the pigsty can be built over the pondsor on the bunds, optimizing space utilization. Cattle andchicken manure is readily available and may be added tofreshwater prawn ponds for fertilization. Goats andsheep may be allowed to graze on the grass of the bunds,thus contributing to efficient facility maintenance. Prawnponds may provide space for ducks to swim; in turn,ducks will fertilize the ponds, control invasive plants,and loosen the pond bottom, releasing nutrients fromthe soil and aerating the water through their swimmingactivities (Tripathi and Sharma, 2001a). These examplesindicate that integrating prawn culture with livestockfarming may be environmentally sound and increaseprofits.

Status of integrated freshwater prawn farmingglobally

The status of a wide range of systems that are currentlyused in the integration of freshwater prawns within pro-duction systems is considered in this section.

Polyculture of prawns with other crustaceans

Martino and Wilson (1986) observed in laboratoryexperiments that the freshwater prawn. M. rosenbergii,and the red swamp crayfish, Procambarus clarkii, showedcompatible behavior to be reared together. Intercroppingwith these species has been described only in the UnitedStates in the 1990s (D’Abramo and Daniels, 1992). Nofurther information is being reported on this topic inaquaculture literature nowadays. The polyculture ofmarine shrimp (Litopenaeus vannamei or Penaeusmonodon) and freshwater prawns is sometimes per-formed in paddyfields or in polyculture with fish in Asia(Azim et al., 2001; Schwantes et al., 2007; Ali et al., 2009;Belton et al., 2009). For example, Azim et al. (2001)reported the existence of mixed culture of M. rosenbergiiand tiger prawns (Penaeus monodon) in brackishwater,but the survival ofM. rosenbergii was low (23–37%). Fur-ther examples of the incorporation of marine shrimp inpolyculture systems in Bangladesh are provided later inthis review.

Fish-prawn polyculture and coculture

A large variety of fish may be reared together with fresh-water prawns in the same ponds. The choice depends onthe region and the availability of fry or fingerlings. New(2002) relates that around the world various freshwaterprawn species are polycultured with single or multiplespecies of fish, including tilapias; common carp (Cypri-nus carpio); Chinese carps—silver carp (Hypophthalmic-thys molitrix), bighead carp (Aristichthys nobilis) andgrass carp (Ctenopharingodon idella); Indian carps—catla (Catla catla), rohu (Labeo rohita) and mrigal (Cir-rhinus mrigala), golden shiners (Notemigonus crysoleu-cas), mullets (Mugilidae), tambaqui (Colossoma spp.),and some ornamental fish. Recently, New and Valenti(2016) extensively examined the scientific literature onthe polyculture of freshwater prawns with tilapia species(mainly Nile tilapia); their conclusions are recorded laterin section “Status of research on integrated freshwaterprawn farming” of this review.

Carp polyculture has been a traditional practice insome Asian countries, mainly Bangladesh and India.This generally involves a combination of the three Indianmajor carps with three or four species of Chinese carps.The introduction of prawns into carp polyculture is apractice that only started to be investigated in the 1980s(Alam et al., 2001; Rahman et al., 2010a). The polycul-ture of carps with freshwater prawns, tilapia and mola(Amblypharyngodon mola, an indigenous phytoplanktongrazing species) has been reported to be more profitablethan traditional carp polyculture or monoculture

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(Hossain and Islam, 2006; Kunda et al., 2008; Rahmanet al., 2010a; Shahin et al., 2011). This is because of thehigher value of freshwater prawns than the fish and indi-cates that the polyculture of carps and freshwater prawnsmay expand in the future.

In Bangladesh, M. rosenbergii is commonly known asgolda. The polyculture of golda with Indian and Chinesecarps has increased in the northern region since the1990s and has popularized freshwater prawn cultureamong existing fish farmers (Hossain and Islam, 2006).Al Noor (2010) has reported that some farmers formerlypracticing freshwater prawn monoculture have started toincorporate four species of native and exotic carps. Thesefarms typically have about 1,500 m¡2 of water surface,which is generally stocked with 20 rohu, 40 silver carp, 5bighead carp, and 2 grass carp (about 200 g each). Thecarps are stocked one or two months after the prawns.Coconut leaves are placed inside the ponds as sheltersfor the growing postlarvae and for prawns to seek refugeduring molting. Commercial feed and farm by-productsare supplied for both fish and prawns. Harvesting occursfrom June to November after 5–6 months stocking. Bythis time, most prawns and carps attain 80–100 g and700–850 g, respectively; total fish and freshwater prawnproduction reaches »250 kg/farm (1.66 t/ha). The majorconstraints are an insufficient supply of PL and the highcost of artificial feed. Species composition and stockingdensities are found to be important factors for maximiz-ing carp production in polyculture with prawns (Islamet al., 2008; Rahman et al., 2010b).

Carps are the most commonly fish used in polyculturewith freshwater prawns in Bangladesh, but other specieshave aroused the interest of producers, such as Thaipanga (Pangasius hypophthalmus) (Islam et al., 2008)and Nile tilapia (Uddin et al., 2006, 2007a, b & c, 2008,2009; Asaduzzaman et al., 2009a & b). Tilapia havegreater market acceptance, are easy to farm, and theirculture cycle is similar to freshwater prawns (Uddinet al., 2007c). The movement of tilapia inside the pondspromotes soil resuspension (Jim�enez-Montealegre et al.,2002). This process may increase benthic dissolved oxy-gen availability, leading to better mineralization andstimulating the natural food web. Asaduzzaman et al.(2009b) concluded that polyculture with tilapia in thepresence of periphyton substrates was more profitablethan prawn monoculture.

In India, the polyculture of M. rosenbergii and M.malcolmsonii with Indian carps started to be practiced inthe 1990s in the northern parts of Karnataka, AndhraPradesh, Orissa, West Bengal, and Punjab, because of thehigher profit achieved when compared to carp or prawnmonoculture (Vasudevappa et al., 2002). Even in fresh-water prawn monoculture, Indian carps such as catla are

sometimes stocked at densities of 100–500 fingerlingsha¡1 to manage excessive algal blooms (Nair and Salin,2006). Nair and Salin (2007) described the use of M.rosenbergii in polyculture of carps in India. The polycul-ture of catla, stocked at 500 fish ha¡1 with M. rosenbergiiis practiced in Andhra Pradesh and other States inpaddy-prawn rotational cropping. Some polyculture sys-tems stock grass carp, catla and rohu in paddy fields at1500 fish ha¡1 and prawns at 2 m¡2. In Kerala, smalland medium size irrigation reservoirs are frequentlyused to produce Indian major carps and M. rosenbergii.Some farmers include bottom-feeding freshwater prawnsin place of common carp and mrigal.

Radheyshyam (2009) reported that in Orissa districts,M. malcolmsonii is reared in polyculture with carps suchas catla (surface feeder), rohu (column feeder), and grasscarp (plant eater). Silver carp can be incorporated intothe species mix when ponds have an abundance of phy-toplankton, but bottom feeder carps such as mrigal andcommon carp are not used in prawn polyculture. Prawnproduction varies according to the level of management,but can reach more than 700 kg ha¡1. In addition to theprawns, 3000–4000 kg ha¡1 of carps can be also pro-duced under semi-intensive management practices.

Extensive polyculture of carps and M. rosenbergii andM. malcolmsonii has been performed in India. Laxmappaand Krishna (2015) reported that stockingM. rosenbergiiPL in the Malampuzha reservoir in Kerala yielded signifi-cant results, with prawn production ranging from 10 to83 kg/ha/year. In addition to the prawns, carps are alsoproduced in the same system. There is no uniformgrowth in freshwater prawns; the harvest sizes rangefrom 40 to 110 g each, due to seed quality and the quan-tity stocked in the reservoirs. The same authors reportedthat extensive polyculture has been practiced in the Koil-sagar reservoir (Telangana State, South India) since2002, with carps such as catla and rohu stocked at verylow densities with prawn juveniles collected from wildsources (M. malcolmsonii) or produced in hatcheries (M.rosenbergii). In this system, water quality, prawn growthand health were not monitored, and supplementary feed-ing and organic fertilizers were not applied. Hence,prawn production was also limited to »10–12 kg/ha/year and fish production was only »75–90 kg/ha/year.In the year 2013–2014, the prawn production of theentire reservoir was of 9.68 t.

New (2002) provided some technical information onpolyculture systems in China involving M. rosenbergii.When the emphasis of polyculture is on the prawns,juveniles of M. rosenbergii are stocked with 1–1.2 cm (at16.5–22.5 m¡2) or 1.5–2.0 cm (at 15–18 m¡2), while big-head and silver carps (12–15 cm) are stocked at 1500–1800 ha¡1. Production of prawns and carps range from

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1500 to 3000 kg ha¡1 per crop and from 750 to 1500 kgha¡1 per crop, respectively. When the emphasis is oncarp production, freshwater prawns are stocked as PL at24–30 m¡2, or as 1.0–1.2 g juveniles at 4.5–9.0 m¡2, oras 1.5–2.0 cm juveniles at 3–6 m¡2, while carps with asize of 3–4 cm are stocked at 16.5–21 m¡2. In this case,the production per ha per crop ranges from 450 to750 kg of prawns averaging 20 g and from 5000 to7500 kg of juvenile carp with 12–15 cm body length.Feed is supplied to the prawns, varying from 20 to 5% ofprawn biomass, being spread around the pond. Sheltersfor the prawns, such as aquatic weeds, grasses, and treebranches are placed in the ponds. Partial seine harvestsare taken and at the final harvest, fish are firstly removedwith a large-mesh seine before the pond is completelydrained. More recently however, Ming (2014) noted thatthe polyculture of prawns with fish in China is a minoractivity and often restricted to experimental research.Personal contacts in 2014 by New & Valenti (2016) indi-cated that there is now little or no activity of this typeeither in China or in Taiwan.

Polyculture is still not common in Brazil but haspotential for expansion, since the low profitability of fishculture causes hundreds of hectares of ponds to beunused all over the country; these could be used for poly-culture (Marques and Moraes-Valenti, 2012) and forincreasing sustainability (Marques et al., 2013). Until2007, polyculture occurred only in the State of Paran�a,southern Brazil, but nowadays occurs in at least fourStates in the South and Southeast regions. In Paran�a,Barreto et al. (2009) stated that the polyculture of tilapiaand prawns (M. rosenbergii) was becoming much morepopular among the tilapia producers. Prawns have ahigher commercial value than fish and thus the eco-nomic return from polyculture was higher than fromtilapia monoculture. Furthermore, some farmers say thatpolyculture improves the quality of water in the ponds.Valenti and Tidwell (2006) reported that ponds arestocked at 2–4 PL or juveniles m¡2. After a week, tilapiajuveniles weighing around 10 g are stocked at 1–3 m¡2.Only tilapias are fed with commercial pellets: the prawnsfeed on benthic organisms and feed residues. Harvestingoccurs after 5–8 months and tilapia productivity is notaffected by the presence of freshwater prawns. Polycul-ture is profitable: the internal rate of return (IRR) wasestimated to be 15–45% and the payback period (PP)3–6 years in the mid-2000s.

It has been reported (E. Ballester, personal communi-cation, 2013) that the producers in Paran�a State werestocking prawns and tilapia at higher densities (6–10prawns and 2–3 Nile tilapia m¡2) and that the best yieldsachieved were 16.6 t of tilapia (average size 500 g) and750 kg of prawns (average size 25 g) per hectare in a

single cycle. The normal productivities achieved are gen-erally »10–12 t of tilapia and 500–600 kg of prawns perhectare per cycle. Currently, 4–8 Nile tilapia m¡2 and 4–10 prawns are sometimes used in Paran�a and S~ao PauloStates (F. Sussel, personal communication, 2015). Besidestilapia, grass carp and bighead carp are also used inprawn polyculture in the States of Santa Catarina andEspirito Santo. In these cases, prawns are the main spe-cies, being stocked at normal densities (10–12 m¡2) andprovided with a pelleted feed, while carps are introducedat 0.2 m¡2 with the aim of controlling invasive weedsand filtering the excess of phytoplankton. Some pro-ducers have substituted carps by tilapia at the same den-sity, since the latter have a better market value.Apparently, tilapia does not compete with prawns forartificial feed at these densities.

Prawn polyculture in Thailand is uncommon, beingpracticed by less than 10% of farmers in mid-2000s(Schwantes et al., 2007). Some prawn producers havestocked tilapia in their systems, but prawns are the maincrop, whereas M. rosenbergii are stocked in tilapia pondsat low densities to provide an additional high value crop(Belton et al., 2009). This practice was a response to thereportedly declining profits of Thai tilapia farmers.

In the Philippines, some farmers polyculture M. rose-nbergii with milkfish, Chanos chanos (Rola, 2007).Prawns (average stocking size of 3 cm) are normallystocked before the milkfish (average stocking size 10 cm)and the average culture period traditionally adopted isfour months for both milkfish and prawn production,with two cycles per year. The average stocking rate formilkfish varies according to the system adopted; how-ever, for prawns the stocking rate is »27,000 ha¡1yr¡1.Polyculture provides additional income for fish farmersbecause of the high market value of prawns. Neverthe-less, some farms believe that the adoption of polycultureneeds financial support. This has constrained the poly-culture of fish and freshwater prawns in the Philippines.

In Egypt, El-Sheriff and Mervat (2009) reported theexistence of commercial polyculture of prawns with Niletilapia. New and Kutty (2010) reported that prawnbroodstock were introduced into Egypt from Malaysiaand Thailand in 1987–1988. Production was said to havepeaked at 10.5 t in 1996 and comprised 7 t from mono-culture, 3 t from polyculture with Nile tilapia, and 0.5 tfrom experimental integrated farming (rice, fish, andprawns). By 1999, the only remaining farm, with a totalarea of 1000 ha, which had 51% government and 49%private ownership and primarily reared mullet, tilapia,sea bream, sea bass, and marine shrimp, was still usingabout 5 ha to rear a small quantity of M. rosenbergii inpolyculture with tilapia and mullet. By 2007, all the pri-vate sector freshwater prawn hatcheries had closed due

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to technical and economic factors and the lack ofdemand for PL. Prawn culture is almost at a standstilldespite promising beginnings and no actual polyculturewith tilapia is taking place (S. Sadek, personal communi-cation, 2014).

There are also reports about commercial or experi-mental prawn polyculture with tilapia in Mexico (Valentiand Tidwell, 2006), USA (Rouse and Stickney, 1982; Tid-well et al., 2000, 2010; Danaher et al., 2007), Puerto Rico(Garc�ıa-P�erez and Alston, 2000; Garc�ıa-P�erez et al.,2000) and Brazil (Valenti and Moraes-Riodades, 2004;Santos and Valenti, 2002; Marques and Moraes-Valenti,2012).

There is a large amount of data on pelagic fish andprawn polyculture that supports the conclusion that nei-ther species affects the other inside the same pond.Figure 5 shows that the growth, survival and yield of tila-pia does not change as prawn density increases. Figure 6shows that the same occurs with prawns as tilapia den-sity increases. In prawn monoculture, on the other hand,both growth and yield are markedly affected by prawndensity (Sampaio and Valenti, 1996; Valenti et al., 2010).Prawn production is related to tilapia production whenonly tilapia are fed (Figure 7). This reinforces the ratio-nale that prawns eat the wastes generated by tilapia.

Commercial fish-prawn polyculture is still relativelyuncommon beyond Asia, despite the substantial science-based knowledge available (Zimmermann et al., 2010).New (2002) suggested that the management of this poly-culture species combination system might be complex,mainly with respect to the lack of synchronicity in har-vesting operations; however, this problem has not beenspecifically reported in the literature. It seems that themajor constraints to the expansion of polyculture in Asiaand Latin America are the same as those for prawnmonoculture, i.e., poor transfer of technology, deficien-cies in PL supply, lack of capital, and undevelopedmarketing.

Rice-prawn culture

Currently, commercial rice-prawn or rice-fish-prawnculture is already commonplace in several Asiancountries, such as Bangladesh, where rice-prawn cul-ture is well developed, playing an important role asan economic and social activity. In other countries, ithas great potential to expand because of the hugeamount of available paddy field areas. In Bangladesh,prawn-rice culture is mainly performed in ghers(Ahmed, 2009; Ahmed et al., 2008b). Barmon et al.,(2004, 2006) reported two main types of gher farm-ing: the brackishwater based shrimp farming per-formed in the coastal region and the freshwater based

rice-prawn farming, performed at the southwesternhigher areas. These include Bagerhat, Khulna, Jessore,Barisal, and Satkhira districts (Ahmed et al., 2008c).In these regions, the practice of small-scale prawnfarming in paddies is widespread due to the availabil-ity of wild freshwater prawn PL, rice fields, a warmclimate, fertile soil, and cheap and abundant labor.Thousands of farmers from that region have con-verted their rice fields to accommodate profitableprawn culture (Ahmed et al., 2010, 2014). In thelower areas of Bagerhat, M. rosenbergii is farmed withthe penaeid shrimp bagda (Penaeus monodon), theIndian carp’s rohu, catla, mrigal, and silver carp. This

Figure 5. Variation of tilapia instantaneous growth rate (IGR), sur-vival, and yield according to prawn stocking density in tilapia-prawn integrated culture around the world. Data points wereobtained from the articles cited in the text. The dashed lines rep-resent the mean values of the dependent variable. r D Pearsoncorrelation coefficient.

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polyculture is performed in fresh to hyposaline water(0–3 salinity), integrating marine shrimp, freshwaterprawns, and fish with low salinity tolerant paddy(BR-23, BRRI-27, 40, and 41) and vegetables rearedon the dike (Ali et al., 2009). In this way, prawn-riceculture allows the economic use of the thousands ofhectares of rice fields that remain waterlogged forfour to six months of the year because of the mon-soon in the upper areas or the intrusion of salinewater in coastal areas; this improves food securityand the socioeconomic conditions of farmers andpromotes the production of rice through biological

pest control and soil fertilization (Ahmed et al.,2010).

Freshwater prawn-rice integrated culture may bemore sustainable system. Barmon et al. (2006) concludedthat in Bangladesh brackishwater shrimp-gher farmingcauses land degradation and has a negative impact onthe environment, livestock, and water quality, whereasthe rice-prawn freshwater gher farming system is friend-lier to the environment, land and water quality, besideshelping to alleviate poverty due to the social inclusion ofmarginal and landless farmers in the system. Ahmedet al. (2012) suggested that rice-fish-prawn farmingmight form part of a “blue-green” revolution to increasefood production in Bangladesh. Coastal aquaculture,which is dominated by export-oriented freshwater prawnand brackishwater shrimp farming (Ahmed, 2013) isthreatened by climate change. Ahmed et al. (2014) pro-posed that integrated prawn-fish-rice farming could berelocated from the coastal region to less vulnerableinland areas, but cautioned that this would requireappropriate adaptation strategies and an enabling insti-tutional environment. Ahmed et al. (2010) reported thatthe annual yield average of prawn in rice fields was467 kg ha¡1, ranging from 387 to 564 kg ha¡1 (small [upto 0.20 ha] and large [above 0.40 ha] farmers, respec-tively). Kamal (2010) reported prawn productivitiesfrom 375 kg ha¡1 in farms larger than 2 ha to 800 kgha¡1 in farms between 0.8 and 2.0 ha. Prawns are fedeither with farm-made feed or commercial pellets, sup-plemented with a variety of other feeds; the preferredtype is the snail Pila globosa, harvested from the flood-plains in large quantities. A range of fish species is com-monly polycultured with the prawns in rice fields, suchas Indian major carps (catla, rohu, and mrigal) andexotic carps (silver carp, grass carp, and common carp)(Ahmed et al., 2008a). The carps are stocked one or twomonths later than prawns, generally at »14,000–20,000prawn PL ha¡1 and 400–570 fingerling carps ha¡1

Figure 7. Relationship between prawn yield (Py) and tilapia yield(Ty). ln D natural logarithm; r2 D coefficient of determination.

Figure 6. Variation of prawn instantaneous growth rate (IGR),survival, and yield according to tilapia stocking density in tilapia-prawn integrated culture around the world. Data points wereobtained from the articles cited in the text. The dashed lines rep-resent the mean values of the dependent variable. r D Pearsoncorrelation coefficient.

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(Kamal, 2010). According to Ahmed et al. (2010), theaverage stocking density of the carps is 2450 fingerlingha¡1 yr¡1 and of the prawns is »18,000–26,000 PL ha¡1

yr¡1. Productivity varies from 387 kg ha¡1 yr¡1 in farmsof up to 0.2 ha to 564 kg ha¡1 year in farms above 0.4 ha;fish productivities are reported as 703 kg ha¡1 yr¡1

(small farms) and 1271 kg ha¡1 yr¡1 (large farms). Cur-rently, the production of prawns aims to supply theinternational market due to their high product value,whereas rice and fish are preferred for householdconsumption.

The yield of rice under gher farming is the same orhigher as the yield from rice monoculture in Bangladesh.According to Mohanty et al. (2004), the increase in riceyield under rice-fish-prawn integration was probably dueto the presence of fish and prawns, since their movementhelps to improve dissolved oxygen levels and stirs up soilnutrients, making them more available for rice. In addi-tion, fish and prawns help to enhance soil organic mat-ter/nutrient status by adding fecal matter and maycontrol plankton population and aquatic insects, in addi-tion to removing bacteria and organic detritus that com-pete with rice for nutrients and energy. This systemtherefore provides a sufficient amount of rice to meetlocal demands and simultaneously produces a high valuespecies (M. rosenbergii) with good market demand. Bar-mon et al. (2004) found that the gher farming systemincreased labor demand compared to rice monoculture.Furthermore, the feasibility of using freshwater prawnsin polyculture with fish allowed the farms to obtain addi-tional fish for home consumption, and also created morejob opportunities for family laborers, benefiting the localpopulation and neighboring communities. Bangladesh isconsidered by Ahmed (2009), as one of the most suitablecountries in the world for integrated prawn-fish-ricefarming, because of its favorable agroclimatic conditions.Consequently, integrated prawn farming has been one ofthe most important sectors of the national economy(Ahmed, 2009; Ahmed et al., 2008b, 2014). Freshwaterprawns are known as “white gold” or “dollar” (Ali et al.,2009), becoming in recent years the most desirable spe-cies in aquaculture (Ahmed et al., 2010; 2014). Ahmedet al., (2014), stated that integrated prawn-fish-rice farm-ing is the most efficient system in terms of resource utili-zation through the complementary use of land andwater. Despite higher production costs per hectare, theaverage annual net return was higher in large farms(US$2,426; farms >0.4 ha), compared to medium(US$1,798; 0.21–0.4 ha) and small (US$1,420; �0. 2 ha)farms (Ahmed et al., 2010). Prawn production in ghersystems has been accompanied by a great deal of socialand economic benefits (Ahmed et al., 2010). Some con-straints to long-term sustainability have been identified,

such as resources, weak transforming structures and pro-cesses, vulnerability context, poor institutional support,and lack of extension services (Ahmed et al., 2008a).Moreover, an insufficient supply of wild PL and diseaseproblems have been identified as major bottlenecks inthe development of the activity (Ahmed and Flaherty,2013; Ahmed et al., 2008c, 2014). To resolve these issues,it is necessary to explore the possibility of developingprawn hatcheries closer to on-growing areas, thus avoid-ing dependence on wild seed and the appearance of dis-eases; this would ensure more sustainable and resilientintegrated prawn-fish-rice culture (Ahmed et al., 2008a,2014; Ahmed and Flaherty, 2013). Despite the con-straints identified, Ahmed and Flaherty (2013) con-cluded that the sustainable development of prawn-fish-rice farming in southeast Bangladesh had great potentialfor improving the food security of farming households,and more broadly the economic growth of the countrythrough earnings from the export of prawns.

In Vietnam, rice-prawn farming is one of the mostimportant models of prawn production. This system iswidespread in the Mekong Delta region. Similarly toBangladesh, Vietnamese rice-prawn farming is per-formed simultaneously (where prawn and rice culturesare carried out together as mixed farming) or in an alter-nate manner (where prawn culture and rice cultivationare carried out on a rotational basis). Productivities varyaccording to the initial size of postlarvae or juveniles,stocking densities, and food management, ranging from40 to 500 kg ha¡1 and from 360 to 900 kg ha¡1, forsimultaneous and alternate systems respectively (Phuonget al., 2006). Hai et al. (2015) stated that prawns are cur-rently stocked at densities of 1–2 m¡2 and production isabout 50–100 kg ha¡1 crop¡1.

Duong (2001) described the main steps that wereemployed in rice-prawn culture in the Mekong Deltaregion 16 years ago. These included the advice that dikesmust be sufficiently high and strong to resist floods;inside the paddies, the depth of water varies from 20 to30 cm. In order to provide shelter during the rearingperiod and to harvest the prawns, peripheral trenches(harvest channels) with 3.0–4.0 m wide and 1.0–1.2 mdepth are excavated. Nets or woven strips of bamboo arefixed at the inlet and outlet water pipes, in order to avoidprawn escapes and to minimize the entrance of preda-tors. Prawn PL and juveniles are collected from the riversand stocked directly in the paddy fields (resulting in lowproduction cost). Residues or by-products of farmingand fishing (cassava, sweet potato, broken rice, soaked orcooked milled rice, rice bran, crabs, and snails) are sup-plied to the prawns. Farm-made feeds consisting of 50%rice bran, 10–20% cooked broken rice, 20–30% trashfish, and 10% oil cake are also used. Fertilization is

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performed with organic and/or inorganic fertilizers andweed control is done manually, without using herbicides.After 4–5 months of rearing, the rice field is drained andthe prawns are collected in the harvest channels.

Phuong et al. (2006) stated that during the dry season inthe Mekong Delta, the salinity of water is quite high, so somepaddies can only be used to rear marine shrimp; however, inthe inland floodplain and during the rainy season, rice-prawnfarming can also be practiced. Trenches usually occupy about20–25% of the paddy area with an average depth of 0.8–1.0 m. Stocking density varies according to the productionsystem: in the integrated rice-prawn culture system, stockingdensity ranges from 1.5 to 5 individuals m¡2 depending onprawn size; in the alternate rice-prawn culture model, thestocking is done with M. rosenbergii PL at 3–12 m¡2 in thetype 1 system, where one prawn crop and one rice crop arecarried out in a year, or with 15 g prawn juveniles (stockingrate not identified) in the type 2 system, where one prawncrop and two rice crops are carried out per year. Productivityranged from»40 to>500 kg ha¡1 in the integrated system,and 900 kg ha¡1 in the alternate culture type 1 and 360 kgha¡1 in the alternate system type 2 systems, respectively. Linand Lee (1992) reported that the yield of prawns in rice inte-gration ranges from 200 to 400 kg ha¡1 in 8 months, with agross annual income of US$ 600–1200.

Halwart and Gupta (2004) described the rearing ofboth M. rosenbergii and M. nipponense in rice fieldsin China. The physical preparation of the rice fieldsis the same as in other countries but thereafter sub-merged aquatic plants are planted in the trenches tocover one-half to one-third of the bottom. For M.rosenbergii, the stocking rate of 1.5 cm sized juvenilesis 3 m¡2, while 4–6 cm sized M. nipponense may bestocked at 3.0–3.8 kg ha¡1 and allowed to breed, orjuveniles are stocked at 23–30 m¡2. Feeding consistsof soybean milk and fish gruel during the early stages(seven to eight days after stocking) and high-proteinpelleted feed or a mixed diet of wheat bran or ricebran with some animal protein source thereafter. M.rosenbergii is harvested before the temperature dropstoo low; meanwhile the harvest of M. nipponense canstart on a selective basis by late November or earlyDecember. The undersized animals are left to growfor the total harvest by May or June, which takesplace before the rice-planting season. Miao and Ge(2002) reported that usually only one crop of prawns(300–450 kg) is produced per year in China, usingsupplementary feeding in addition to the normal riceproduction. These authors pointed out that it is avery effective approach to improve the economicreturn from traditional rice cultivation and also has avery sound environmental effect, due to the signifi-cantly reduced use of pesticides and other chemicals.

In northeastern India, Das (2002) stated that an indig-enous traditional capture fishery system in rice fields pre-vailed at that time. Due to the gradual reduction ofnatural stocking that they observed, these authors rec-ommended that the traditional wild capture systemshould be improved or replaced by adopting suitable cul-ture techniques that would increase yields, enhancingsocioeconomic development in rural areas. In Kuttanad,State of Kerala, Kurup and Ranjeet (2002) reported that,as occurs in Bangladesh, the simultaneous culture of riceand prawns was substituted by the rotational farming ofrice and M. rosenbergii. This occurred mainly becausethe double cropping of rice may not always be feasibledue to flooding during the monsoon season and to thehigh cost of land and labor. Rotational culture is notonly helpful in improving farming revenue, but also pro-vides additional employment. This practice is beneficialbecause it reduces the cost of pond fertilization, main-tains soil fertility, avoids the accumulation of waste prod-ucts, improves pest control, and enables farmers tocontinue with their traditional forms of livelihood.Prawns are stocked from 1.5 to 6.0 m¡2 in rice-prawnmonoculture and from 0.2 to 1.5 m¡2 in rice/prawn/fishpolyculture (0.1–0.5 fish m¡2). The productivity inmonoculture ofM. rosenbergii ranges from 95 to 1300 kgha¡1 while between 70 and 500 kg ha¡1 of prawns and200 and 1200 kg ha¡1 of fish (mainly catla, rohu, andgrass carp) are obtained in polyculture. Some integratedrice-prawn farms in Kuttanad that do not use chemicalfertilizers or pesticides have been certified as “organicfarms” (Nair and Salin, 2009; Nair el al., 2014). Theseauthors reported that rice productivity decreased about23% under organic farming; however, the organic prawncrop yield of 396 kg ha¡1 was 10% higher than the yieldof the conventional system (360 kg ha¡1). Total invest-ment for organic rice and prawns was approximately 20and 17% greater than for the non-organic products.These higher costs were compensated for by the productvalues, which were 36 and 26% greater than conventionalproducts, respectively. Organic rice farming culturebrings lower returns than the conventional system. Thecombination of organic rice farming with organic prawnfarming enhanced net revenue by 20% over conventionalrice/conventional prawn production (Nair et al., 2014).

Halwart and Gupta (2004) stated that rice-fish farm-ers in East Java (Indonesia) useM. rosenbergii in polycul-ture with milkfish (Chanos chanos) and silver barb(Barbodes gonionotus), either in concurrent or rotationalsystems. According to Giap et al. (2005), rice-prawn cul-ture in Thailand is extensive, based on natural food.These authors reported that the profits were 43–91%higher in rice-prawn culture than in rice monoculture,probably because of the higher value of prawns

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compared to the fish species usually cultured in ricefields. Schwantes et al. 2007 noted that although themonoculture of M. rosenbergii was well developed inThailand, crop rotation and integrated culture were rareor non-existent, despite their huge potential.

Constraints to the expansion of rice-prawn culture inAsian countries have been presented by several authors.According to Giap et al. (2005) the increasing use of pes-ticides and chemicals in rice production is the main bot-tleneck. Phuong et al. (2006) considered that there is adanger that unit price would decrease with increasedproduction and detrimental effects on the local environ-ment would arise. Ahmed and Garnett (2010, 2011) andAhmed et al. (2010) reported a number of constraintsfor rice-fish-prawn farming in Bangladesh, including ashortage of operating capital, increased production costs,lack of prawn PL due to the decrease in natural stocks,an inadequate supply of snails for feed, prawn diseases,climatic disasters (flood and drought), and social con-flicts. Nabi (2008) highlighted the fact that rice-prawntechnology is still not structured to address the foodsecurity goal of smallholders and therefore is adoptedonly by those who are better placed to assume the risksof the activity. Kamal (2010) included as constraints thehigh costs of prawn PL and supplementary feed. Partialsolutions for these constraints would be the expansion ofprawn hatcheries and the use of low-cost locally pro-duced ingredients for feed other than snails; theseimprovements would help to increase production, reducenegative environmental impacts, and increase job oppor-tunities. Recent climate change has also been considereda risk to growth in this activity (Ahmed et al., 2014).Boock et al. (2013) considered that the low level of waterin Brazilian paddies may causes large temperature oscil-lations and possible prawn mortality, and that the onlyway to prevent this would be to restrict the period ofrearing to the warmer months of the year.

In continents other than Asia commercial rice-prawnssystems have not been developed but the crawfish Pro-cambarus clarkii has been reared in small- and medium-scale in the Southeast of the United States, integrated withrice or soybean production in alternate systems (McClainet al., 2007). The economic importance of this system inthat country is far from that existing in Asia. Someresearch on rice-prawn farming outside Asia will be pre-sented in section “Status of research on integrated fresh-water prawn farming” of our review.

Other integrated systems

There are some commercial initiatives involving the inte-grated culture of freshwater prawns. The use of pondmargins for planting bananas, coconut, and papaya is

common in Asian countries and also in the state ofEspirito Santo (Brazil). The use of the effluent from fishand prawn ponds to irrigate the cultivation of greens,fruits, and vegetables occurs, mostly in Asian countries,but this practice is increasing in Europe and NorthAmerica (Diver and Rinehart, 2010). Lin and Lee (1992)reported the existence of vegetable-prawn integrated cul-ture in areas converted from rice fields. This system con-sisted of ditches for prawn rearing and platforms andembankments for growing cucumbers, string beans, egg-plants, and bananas, thus providing supplementaryincome as well as food for domestic consumption.Prawns were stocked at 4–6 m¡2 and fed twice daily amash of rice bran, cassava, and trash fish. Each 6 months,about 100 kg of prawns were cropped. Ahmed et al.(2008a) stated that almost all gher farmers in Bangladeshcultivate dike crops mainly for household consumption,while 32% of farmers sold their crops in local markets.During the winter season, different types of dike cropssuch as carrots, tomatoes, onions, mustard, and “yard-long” beans were produced, while crops produced in thesummer season included ladies’ fingers, sweet gourdsand other vegetables. Most of the farmers (88%) pro-duced more than three dike crops in different seasons,while the rest of the farmers produced three or less.Fruits, such as bananas, papayas, and guavas are also cul-tivated. Also in Bangladesh, Ali et al. (2009) studied sixcommercial ghers (9.08 ha total area) and reported thatthe total production of vegetables cultured along thedikes of rice fields were: papayas (1,140 kg), eggplants(750 kg), bitter melons (400 kg), ladies’ fingers (521 kg),tomatoes (588 kg), peppers (108 kg), bottle gourds(998 kg), and pumpkins (1248 kg); these produced a totalincome of 3808 TK (US$ 48.70) per hectare per cycle.

Aquaponic systems generally use fish species that adaptto intensive recirculated aquaculture systems (RAS),including tilapia, trout, and perch. Most commercial aqua-ponics systems in North America are based on tilapia, butin Australia barramundi (Lates calcarifer) is the species ofchoice (Diver and Rinehart, 2010). As freshwater prawnsare rarely reared in RAS, their inclusion in aquaponic sys-tems is still very restricted. There are records of aqua-ponics systems that include tilapia in polyculture with M.rosenbergii (Martan, 2008). Some aquaponics producersrecommend rearing freshwater prawns at low densitieswithin hydroponics tanks for pest control, waste disposaland as a source of extra income (Friendly Aquaponics,2013; Somerville et al., 2014).

The integration of fish-prawn polyculture with therearing of ducks, chicken, and pigs has been reported byTripathi and Sharma (2001a, b) in some Asian countries,mainly in India and Vietnam, but occurs only in family-run systems.

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Status of research on integrated freshwaterprawn farming

This section reviews the current published researchinformation on integrated freshwater prawn farming.

Fish-prawn polyculture

Zimmermann et al. (2010) presented an extensive reviewof research on the polyculture of prawns with variousspecies of fish. This review included the aspects of man-agement, water quality, feeding, stocking strategies, andprofitability. This publication should be considered bythose who seek more details about these topics.

More recently, New and Valenti (2016) reviewed theliterature on the specific polyculture of freshwaterprawns with tilapias, and noted that a considerableamount of research on tilapia-prawn polyculture hadbeen performed during the past four decades. Mostpapers on this topic concentrated on the giant riverprawn (M. rosenbergii) and Nile tilapia (O. niloticus).The problem is that the available information is frag-mented and many important subjects are not covered atall. It is almost certain that prawns do not affect the per-formance of tilapia in earthen ponds and stocking themin such ponds may significantly enhance total revenues.According to New and Valenti (2016), polyculture is rec-ommended for tilapia-pond culture farmers, who wantto retain tilapia as the major species. On the other hand,the presence of tilapia in prawn ponds may reduce prawnyield, but is unlikely to reduce profit.

The main constraint is the feeding of prawns whenpolycultured with tilapia (i.e., free-swimming within theponds) (New and Valenti, 2016). Tilapia consumes pel-leted feed very rapidly and eats much more than theyneed for growth (superfluous feeding). Thus, prawns donot have the opportunity to ingest the supplied pellets.In addition, supplying sinking pellets may attract tilapiato the pond bottom, which may create negative interac-tions with the prawns. Thus, it is advisable for those whowant to intensively feed prawns in order to obtain highprawn productivity to use the coculture technique (i.e.,stocking tilapia in cages within prawn ponds). It seemsthat caged tilapia can improve water quality by filteringalgae, mainly cyanobacteria, and provide residues, likefeces and waste feed, which can be ingested by prawns,thus reducing the quantity of commercial diet needed. Itis important to consider the local costs of cages. Whencontemplating the establishment of new tilapia-prawnfarms, the comparative economics of rearing tilapia inpolyculture with prawns compared to coculture needs tobe carefully assessed.

The stocking size in earthen ponds of both tilapia andprawns in polyculture seems to be irrelevant, since tilapia

are stocked after the fry phase, generally at < 3 g (Newand Valenti, 2016). No injuries inflicted by prawns ontilapia have been described. Reduction in prawn survival,when it occurs, is more likely to be caused by the inade-quacy of the feed available to the prawns than by anydirect aggressive action of the tilapia (except that of tila-pia fry on prawn postlarvae). The use of male sex-reversed tilapia is highly recommended in polyculture. Ifthey are not available, it is better to coculture the tilapiain cages to avoid reproduction (fertilized eggs will fallthrough the cage mesh and cannot be incubated in themouth of the female).

Stocking density and the proportion of tilapia andprawns stocked appear to be important factors (New andValenti 2016). The optima certainly depend on the sitecharacteristics (such as water quality and natural produc-tivity), the general management system utilized (such asthe use of artificial substrates and aerators), and thechoice of the major species (tilapia versus prawns), whichwill be determined by the characteristics of the intendedmarket. Thus, it is difficult to generalize; the informationavailable is not sufficient to show a general picture. Mostresearch has been performed using 1–2 tilapia m¡2 and2–4 prawns m¡2. It is the total biomass of both speciesrather than the individual numbers that should be con-sidered; this factor has been neglected by investigators.The best situation would be to consider the amount ofwaste feed necessary for the prawns and to provide onlythe tilapia with floating pelleted feed. In this way, a realmulti-trophic and multi-spatial equilibrated system(IMTA) could be achieved. To obtain a sustainableIMTA, information on natural biota and trophic chainswithin ponds, energy and nutrient balances, and themanagement necessary to drive energy and key nutrients(N, P, and C) to the target species should be determined.These are important avenues for new research and couldallow the use of various natural foods within the ponds,thus decreasing the supply of allochthonous feed. Thiscan also increase the environmental and economic sus-tainability of the systems.

During the seventies and the eighties, some trials wereconducted in Israel and in the United States, focusing onthe polyculture of M. rosenbergii with a mixture of fishspecies in ponds frequently enriched with manure (Buckand Bauer, 1980; Malecha et al., 1981; Buck et al., 1981,1983; Costa-Pierce et al., 1984, 1987; Wohlfarth et al.,1985). The most common species of fish were carps (sil-ver carp, big head carp, grass carp, and the commoncarp) and tilapias. In research work carried out in theUnited States, production of prawns varied greatly. Mal-echa et al. (1981) reported averages that ranged from 264to 414 kg ha¡1 and Wohlfarth et al. (1985) obtained 66to 791 kg ha¡1 of prawns. This huge variation in

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production may occur due to the feed management sys-tems adopted, based on water fertilization with manurethat can differentially influence the primary productionin the ponds, as a function of diverse ecological interac-tions. In other research, Costa-Pierce et al. (1984) relatedthat the polyculture of carps (silver and grass) with M.rosenbergii combined with reduced feeding rates may bea simple solution to the pervasive problems of waterquality control and benthic reducing conditions. Martinoand Wilson (1986) noted that the interactions observedin the polyculture of M. rosenbergii, Mossambica tilapiaand the crawfish Procambarus clarkii were intraspecificand that the cannibalism observed among the crusta-ceans was not influenced by the presence of tilapia. AlsoCosta-Pierce et al. (1987) observed that the polycultureof M. rosenbergii with silver carp, grass carp and greymullet (Mugil cephalus) resulted in a total production ofall species twice as high than in prawn monoculture,even at a low feeding rate and despite the existence ofcompetition for resources among the species inpolyculture.

Also in the United States, Rouse et al. (1987) foundthat optimum prawn survival (93.8%) was attained atlow stocking densities (4 m¡2) in polyculture and thattilapia reproduction had a negative impact on prawngrowth. Scott et al. (1988) studied the polyculture of M.rosenbergii with the golden shiner Notemigonus crysoleu-cas, a cyprinid fish largely used as a bait fish, and foundthat prawn yield was significantly higher in polyculturethan in monoculture. On the other hand, shiner survivalwas significantly higher in monoculture and shiner yielddid not differ between monoculture and polyculture.

In Israel, the polyculture of prawns with several finfishspecies, including carnivores, was commonplace in the1980s and the success of the stocking strategy was foundto depend on the use of compatible stocking sizes of eachspecies (Hulata et al., 1988). These authors recom-mended the introduction of carps and tilapias withprawn juveniles weighing 0.25–0.50 g at a density of2 m¡2. This stocking density was also used by Karpluset al. (1987), who obtained prawn survival up to 87%and yield among 332–560 kg ha¡1 rearing juveniles ofvarious sizes in 400 m¡2 ponds in polyculture with tila-pias (Oreochromis sp.), common carp (Cyprinus carpio),silver carp, and grass carp. Cohen and Ra’Anan (1983)concluded that an increase in tilapia density had noinfluence on prawn yields or on the social structure ofM. rosenbergii reared in polyculture with tilapia. Prawnyields and average weights were affected by the densityof the prawn population alone, and survival rates in allcases were above 85% and unrelated with either prawnor tilapia stocking rates. On the other hand, theseauthors found that the growth of tilapia when grown

with freshwater prawns was strongly affected by its ownstocking density and by the feeding-manuring strategy.Karplus et al. (1987) stocked three fractions of size-graded juveniles of M. rosenbergii in polyculture withtilapia and carps (silver, common and grass) andobserved that the fraction of “upper” prawns (32% of thetotal) resulted in a smaller proportion (8%) of smallmales and higher proportion of blue-claws (22%), poten-tially providing a net income almost nine times that ofthe control. Mires (1987) did not find significant differ-ences in yield when stocking M. rosenbergii juveniles at5–7.5 m¡2 in polyculture with Nile tilapia fingerlings at0.6–0.7 m¡2. Hulata et al. (1990) found that there was nosignificant effect caused by the age or size of juvenile M.rosenbergii stocked on the production of prawns in poly-culture with tilapias (hybrid and red) and carps (com-mon, grass and silver).

In Bangladesh, Haque et al. (2003) determined theoptimum density of M. rosenbergii in polyculture withcatla, rohu and mrigal to be 6000 ha¡1. In Pakistan, Mia(2004) studied the effect of stocking prawns at three den-sities (6000, 8000, and 10,000 ha¡1) with a single fishdensity of 5000 ha¡1, using several carp species, such assilver carp (35%), catla (15%), rohu (30%), and mrigal(20%) in one experiment and silver carp (30%), catla(15%), rohu (34%), mrigal (5%), grass carp (15%), andblack carp (1%) in a second experiment. The best pro-duction achieved was 122 kg ha¡1 of prawns and4200 kg ha¡1 of fish, corresponding to the initial prawnstocking density of 6000 ha¡1 and four carp species inthe first experiment. It is notable that the temperaturedropped below 10�C during the experiments, which cer-tainly would have caused high prawn mortality. Simi-larly, Hossain and Islam (2006) tested five stockingdensities ofM. rosenbergii (2500, 5000, 7500, 10,000, and12,500 ha¡1) in polyculture with catla, rohu and silvercarp at fish densities of 2500, 5000, and 2500 ha¡1,respectively, during three months of culture in tempera-tures ranging from 27 to 31�C. Prawn survival washigher than 75% and fish »90%. According to theseauthors, the overall total production of prawns and fishwas significantly higher at prawn stocking densities of7500 and 10,000 ha¡1 (2916 and 2914 kg ha¡1, respec-tively) and the highest profit was obtained at a prawndensity of 10,000 ha¡1. Prawn production increased withdensity and varied from 86 to 361 kg ha¡1. Hossain andKibria (2006) studied the polyculture of M. rosenbergiiwith rohu and catla in ponds during the winter using for-mulated diets, obtaining a production of prawns rangingbetween 253.1 and 323.8 kg ha¡1 in the five-month cul-ture period. These authors concluded that the overwin-tering rearing of freshwater prawn/fish polyculture isfeasible; however, further study was thought to be needed

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to determine the optimum stocking density of M. rose-nbergii and carp polyculture, for these specificconditions.

Jana et al. (2007) found a significant reduction inmean counts of heterotrophic bacteria, ammonifyingbacteria, protein mineralizing bacteria, and nitrifyingbacteria in prawn-carp polyculture when compared toprawn monoculture or mixed-carp culture; this indi-cated that there was less accumulation of uneatenfeed in the polyculture system when prawns werepresent. Siddique et al. (2010) studied the abundanceof zooplankton and growth of M. rosenbergii in poly-culture with silver carp and catla, concluding thatstocking prawns at 1.5 m¡2 resulted in slightly highertotal productivity (fish plus prawns) than stockingprawns at 1.0 m¡2 for identical fish densities. Jahan(2011) tested the effects of two types of feed (100%rice bran and a mixture of 20% rice bran, 30% fishmeal, and 50% oil cake) on the production of prawnsand carp. Prawn productivity varied from 798 to1089 kg ha¡1 and the best results were achieved forthe mixed diet. Jasmine et al. (2011) shewed that thepolyculture of M. rosenbergii, catla and silver carpwas more economically feasible than the polycultureof carps alone (catla, rohu, mrigal, and silver carp).Pervin et al. (2012) reported that the inclusion ofmola at 2 m¡2 in polyculture with M. rosenbergii didnot affect the production performance and survival ofprawns stocked at 3 juveniles m¡2. Ahsan et al.(2013) concluded that the introduction of rohu at adensity of 500 ha¡1 increased the net profit and thecombined production of prawns and finfish in poly-culture systems of M. rosenbergii, silver carp, catla,and mola.

In Bangladesh, Rahman et al. (2010a) stated thatprawn juveniles were expensive inputs (about 47% of thetotal cost) in all treatments, followed by prawn feed (16–17%). About 70–77% of the total income was obtainedfrom the proceeds of prawn sales, despite the smallerbiomass of prawn production (31–37% of the total). Inanother trial, Rahman et al. (2010b) studying the poly-culture of silver carp, catla, and all-male freshwaterprawns, concluded that although the prawn biomass(38%) was smaller than the fish (62%), its value washigher, resulting in about 76% of total benefit. Similarresults were obtained by Hossain and Islam (2006): thehighest profit was obtained in the treatment with aprawn stocking density of 10,000 ha¡1, probably due tothe higher production of prawns and their high sale val-ues. Alam and Murshed-e-Jahan (2008), evaluating thetechnical and cost efficiency of prawn-carp systems of105 farmers of Bangladesh, reported that 50% of farmersdisplayed full technical efficiency, but only 9% were cost

efficient, concluding that technical efficiency may onlybe a short-term concern and the farmers have to be eco-nomically efficient in the long term as well.

Other technologies have been investigated and associ-ated with the production of prawns in polyculture sys-tems. Rahman et al. (2010b) determined the bestdensities of silver carp (1500 ha¡1) and catla (1000 ha¡1)in polyculture with mola and all-male M. rosenbergii.Also Rahman et al. (2010a) found that selective prawnharvesting had a more beneficial effect than the ablationof the chelae of M. rosenbergii males on prawn and fish(silver carp, catla, and mola) productivity. Kunda et al.(2009b) found that all-male M. rosenbergii polyculturewith catla, rohu, and mola was economically more viablethan all-female and mixed-sex prawn polycultures.Kunda et al. (2009a) studied the effects of including catla(Catla catla) and Nile tilapia in a prawn-mola polycul-ture performed in rice fields after the harvesting of rice(rotational rice-fish culture system). Mola (»2 g) plusprawns (»3 g) were each stocked at 2 m¡2. The addi-tional stocking of either tilapia (»10 g) or catla (»43 g)at 0.25 m¡2 increased total crop and profit. Theseauthors also reported that the introduction of catlaresulted in higher cost: benefit ratio and higher com-bined production of fish and prawns than the introduc-tion of tilapia or tilapia plus catla in the system. Tilapiadid not affect prawn size, survival, or productivity, butsignificantly reduced the zooplankton and phytoplank-ton concentration in the water column, creating a betterwater condition for both fish and prawns.

The use of biofloc technology in M. rosenbergii-carppolyculture was studied by Prajith (2011). This authorconcluded that 25% rohu and 75% catla should beincluded in the system, since these species equally havethe ability to harvest the biofloc. Catla consumes theplankton enhanced by the floc whereas rohu graze onthe bacterial protein.

The periphyton-based system is a promising tech-nique that has shown benefits for tilapia (Oreochromisspp.) and M. rosenbergii in polyculture. In Bangladesh,Uddin et al. (2007b) found that the introduction of sub-strates (bamboo poles) improved survival, final weightgain and net yield of both tilapia and prawns. In thisexperiment, 1.8 g fingerlings of Nile tilapia were stockedat 2 m¡2 and 30-day juvenile prawns at 2 m¡2 during145 days in experimental ponds. Substrate additionresulted in a 46% higher production of tilapia, whileprawn production increased by 127%. The authors alsoconcluded that prawns did not affect tilapia performanceand that there is a very low dietary overlap between thetwo species. Uddin et al. (2006) studied the polyculturein a substrate-based system of genetically improvedfarmed tilapia (GIFT) and prawns, determining that the

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best stocking ratio was 75% GIFT and 25% prawns, interms of productivity and economic return. Uddin et al.(2007a, c), examining the effect of three total stockingdensities—20, 30, and 40 m¡2, (each at a 3:1 tilapia:prawn ratio)—found that 30 m¡2 produced a the bestnet yield (2209 kg ha¡1 of tilapia and 163 kg ha¡1 ofprawns) and cost-benefit ratio.

Uddin et al. (2008), studying the combined effects ofperiphyton, tilapia, and prawn stocking densities andfeed on pond water quality, concluded that the use ofsubstrates for periphyton growth was a low-cost culturemethod that results in a more favorable environment forthe cultured organisms. This resulted from the avoidanceof organic loading and the simultaneous provision of anextra source of food. In addition, the synergistic relation-ships between tilapia and prawns, through their effectson pond ecology and the good growth rates obtained atrelatively high stocking densities of both organisms, indi-cated that their polyculture was technically feasible andeconomically viable. Uddin et al. (2009) found a favor-able 50–57% net profit margin in trials conducted toinvestigate the effects of substrate addition and supple-mental feeding on plankton composition and productionin tilapia and prawn polyculture. The net return washigher for substrate-based ponds than for control ponds,indicating that it is profitable to use bamboo in this way.Asaduzzaman et al. (2009a) determined the optimumstocking rates for each species from the point of view ofeconomic return. These stocking rates (5 g prawnsstocked at 3 m¡2 and 24 g tilapia stocked at 0.5 m¡2)were used by Asaduzzaman et al. (2009b) in a furthertrial. The addition of periphyton substrates (bambooside shoots) in C/N controlled ponds (C/N D 20:1) sig-nificantly benefited the survival and yield of prawns.

In India, Reddy et al. (1988) found that the polycul-ture of M. malcolmsonii with the Indian carps rohu,catla, and a hybrid catla £ rohu gave a significantlyhigher prawn survival rate (51.4–68.8%) than when com-mon and grass carps were the fish species stocked (6.8%prawn survival). John et al. (1995) found that both M.rosenbergii andM. malcolmsonii performed better in pol-yculture with common and silver carps than in monocul-ture or in bispecies culture. Productivity of both prawnsand fish shows great variability under polyculture, asmight be expected since the trials were conducted underdifferent conditions. Vasudevappa et al. (2002) found aproduction of 2418 kg of catla plus rohu and 780 kg ofM. rosenbergii per hectare, with survival rates of 98 and80%, respectively. Singh (2003), polyculturing M. rose-nbergii with Indian major carps (catla, rohu, and mrigal)in saline land holdings of the southwestern districts ofPunjab in North India, obtained 2655 kg ha¡1 of carpstogether with 150 kg ha¡1 of prawns with a mean weight

of 150 g for males and 50 g for females. Mohapatra et al.(2007) found that the polyculture of carps with M. rose-nbergii indicated that higher production levels of 202%for fish and 156% for prawn could be achieved in green-house ponds as compared with outdoor ponds. Soundar-apandian et al. (2008) found that the best stockingdensities for the polyculture of M. rosenbergii(25,000 ha¡1) and Indian carps (10,000 ha¡1) resulted inthe production of 381 kg ha¡1 of prawns and 500 kgha¡1 of fish.

The polyculture of M. rosenbergii with channel catfishIctalurus punctatus has been experimentally studied inthe United States. Huner et al. (1983) reported that whenprawns were stocked at 25,000 ha¡1 with catfish fry at150,000 ha¡1 in nursery ponds where there was also aresident crayfish population, both prawn production andsurvival were low (92–270 kg ha¡1 and 15–22%). Miltneret al. (1983) stocked nursed 0.5–1.0 g prawns at a lowdensity (2500 ha¡1) with channel catfish, either finger-lings (11–12 g) at 3700 or 7400 ha¡1, fry (0.05 g) at100,000 or 200,000 ha¡1. All ponds also contained 3–4 gsilver carp at 740 ha¡1 and 250–300 g white amur (grasscarp) at 100 ha¡1. Prawn yields were not significantlydifferent between the various catfish treatments andranged from 156 to 170 kg ha¡1, compared to 200 kgha¡1 in the control (prawns, no catfish). Differences inprawn survival among the various catfish treatmentswere also not significant (93.3–98.8%). Pavel et al. (1985)confirmed the better performance of prawns stocked atlow density (4940 ha¡1), reporting yields of 124–125 kgha¡1 and survival of 76–85% in grow-out ponds duringspring. D’Abramo et al. (1986) stocked juvenile prawns(0.1–2.6 g) at 4942 ha¡1 in polyculture with fry, finger-lings and adult catfish and fed only the fish, achieving aprawn yield of 172 kg ha¡1 and a survival rate of 93%.Heinen et al. (1987) found that a mean survival rate andyield of 75% and 152 kg ha¡1, respectively, could beattained at similarly low prawn stocking densities(0.5 m¡2) in catfish fingerling ponds. Lilyestrom et al.(1987) found that, in polyculture with catfish, M. rose-nbergii fed mainly on insects, macrophytes and the cat-fish diet. The contribution of formulated feed to prawngrowth increases as animals grow. Apparently, polycul-ture has no adverse effect on the survival, growth, or pro-duction of either species. Commercial polyculture offreshwater prawns with fish is almost non-existent in theUnited States, despite the fact that these promisingresults were obtained. Tucker et al. (2004) stated thatcommercial polyculture with adult catfish is impractica-ble due to the difficulty in harvesting both species simul-taneously, but on the other hand proposed thatpolyculture with fingerlings could be a good alternative,since harvesting could be carried out with a seine with a

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2.5 cm2 mesh, that selects the prawns and allows the cat-fish fingerlings to escape.

Studies on the coculture of Nile tilapia in cages sus-pended in ponds stocked with M. rosenbergii have beencarried out in the state of Kentucky. Tidwell et al. (2000)found that the use of distillers’ grains with solubles as adirect feed for the tilapia was economically more inter-esting than the use of commercial feed. Furthermore, theaddition of tilapia in polyculture increased total pondproductivity by approximately 81%. Danaher et al.(2007) studied the effects of various densities of cagedNile tilapia on water quality, phytoplankton populations,and prawn and total pond production in ponds stockedwith M. rosenbergii at 69,000 prawns ha¡1. This experi-ment consisted of three treatments (prawn monoculture,low-density coculture with two cages of 1 m3 (100 fishper cage) and high-density coculture with four similartilapia cages. The total culture period was 106 days fortilapia and 114 days for prawns. The overall mean after-noon pH level and phytoplankton biovolume were sig-nificantly lower in coculture ponds than in monoculture.Prawn weights were significantly higher in coculturethan in monoculture, but the survival of both prawnsand tilapia did not differ between treatments. Prawn pro-duction was significantly greater in high-density(2720 kg ha¡1) than in low-density coculture (2368 kgha¡1), which in turn was greater than in monoculture(2125 kg ha-1). The prawn feed conversion ratios (FCRs)were significantly lower than in monoculture, while thespecific growth rate (g/d) were significantly higher incoculture ponds. There were no significant differences inthe percentage of marketable prawns (>20 g) betweentreatments, with an overall average of 93.3%; however,both coculture treatments had a significantly higher per-centage (83.0%) of premium prawns (>30 g) than mono-culture ponds (70.4%).

The effects of confined (coculture) and unconfined(polyculture) tilapia with freshwater prawns on prawngrowth, tilapia growth, algal populations, and water qual-ity were studied by Tidwell et al. (2010). Juvenile prawnswere stocked at 62,000 ha¡1 in nine ponds. Three controlponds contained only prawns; monosex (male) Nile tila-pia were stocked at 4400 ha¡1 unconfined in three otherponds; and the same size and number of tilapia werestocked but confined in two, 1 m3 cages at 100 fishcage¡1 in three additional ponds. Prawns and tilapiawere fed with commercial pellets. In the polyculturetreatments, average prawn harvest weight (27 g) andprawn production (1625 kg ha¡1) were significantlylower and prawn FCR (3.0) was significantly higher thanin the other two treatments. There were no significantdifferences (P > 0.05) between the monocultured andthe cocultured treatments in terms of prawn harvest

weight, production and FCR, with combined averages of38 g, 2465 kg ha¡1, and 1.9, respectively. Tilapia in cocul-ture had a significantly higher survival rate (99.7%) andFCR (1.5) than in polyculture (90.3 and 0.8, respectively).There were no consistent trends in treatment differencesamong water quality variables or phytoplankton popula-tions. These authors stated that the lower prawn produc-tion in the polyculture treatment is probably due tocompetition for food with the large number of tilapiajuveniles, since despite the use of monosex males tilapia,several cohorts of juvenile tilapia were produced in poly-culture ponds, resulting in over 2500 kg ha¡1 of juve-niles. These authors concluded that the confinement oftilapia in cages (coculture) appears preferable to uncon-fined culture of tilapia (polyculture) with freshwaterprawns.

Studies on the rotational polyculture of M. rosenbergiiand red swamp crawfish Procambarus clarkii were car-ried out in the United States about 25 years ago. Grana-dos et al. (1991) obtained 157–248 kg ha¡1 of 11–17 gprawns and 746–1266 kg ha¡1 of crawfish when prawnswere stocked at 17,500 ha¡1. Brood crawfish werestocked in May when they borrowed into the pond bot-tom. In July, prawn PL were stocked and draining of thepond and prawn harvesting occurred in October. Thepond was refilled and crawfish were harvested from Jan-uary to May by trapping. Prawns were of a good size forsoft shell production during the months when crawfishwere not available for soft shell production. Avault(1990) and Caffey et al. (1993) mentioned that the pro-duction of soft-shell M. rosenbergii was viable in autumnand soft-shell crawfish in summer using the same facili-ties. D’Abramo and Daniels (1992) studied the same sys-tem and obtained a better production of 1444 kg ofcrawfish and 855 kg ha¡1 of prawns, concluding that thesystem was economically feasible.

Some studies on prawn polyculture were carried outin Brazil during the last 25 years. Mendes et al. (1998a)polycultured M. jelksii with two ornamental fish, Ptero-phyllum scalare and Poecilia reticulada over a period oftwo months; the survival rates of the fish were 84 and92.5%, respectively, while prawn survival was 49%.Mendes et al. (1998b) reared Xiphophorus helleri (also anornamental fish) in nursery tanks of M. rosenbergii fortwo months; mean survival was 60.5 and 63.5%, respec-tively. Silva et al. (2008) reported that the polyculture ofornamental goldfish (Carassius auratus) and angelfish(Pterophylum scalare) withM. rosenbergii in 50 L aquariaduring the nursery phase, using the densities of 20 M.rosenbergii PL, 8 P. scalare, and 1 C. auratus per aquar-ium was feasible.

Experiments on the polyculture of freshwater prawnswith food fish have also been carried out in Brazil.

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Carvalho et al. (1998) found that the polyculture of M.rosenbergii with common carp (Cyprinus carpio) inNortheastern Brazil was more profitable than eitherprawn or carp monoculture. Production was 606 kg/ha/4.5 months and 2050 kg/ha/4.5 months, respectively, butprawns accounted for 62% of the total income. Ara�ujoand Portz (1997) compared two systems of polyculture(tilapia-prawn and common carp-prawn) in three differ-ent stocking densities (0.5–0.5, 0.75–0.75, and 1.5–1.5 m¡2) and found that the polyculture with tilapia at1.5–1.5 m¡2 provided the best productivity (4240 and842 kg ha¡1 of tilapia and prawn, respectively), probablydue to the minor competition for feed with tilapia thanwith carps. Prawn densities and the economic feasibilityof tilapia and prawn polyculture was studied by Santos(2001) and Santos and Valenti (2002), who tested threeprawn densities (2, 4, and 6 PL m¡2) with one fish den-sity (1 fish m¡2). The density of 4 m¡2 resulted in higherproductivity (909 kg ha¡1) and a mean weight of 23.0 gand survival of 92% in a cycle of 175 days. The IRRranged from 15 to 45%, while the PP was of 2.5–6 years,depending on the selling price of tilapia. These authorsalso concluded that stocking densities of up to 6 prawnsm¡2 did not affect tilapia production and the rearing sys-tem required neither additional feeding nor significantchanges in management. Tilapia monoculture showedno profitability, while the polyculture system allowed anincrease in total production with the same amount ofsupplied feed.

The polyculture of tilapia fingerlings (33 m¡2) duringthe nursery phase and Amazon river prawns (14 m¡2)was studied in a commercial farm by Boock et al. (2008),using a 1500 m2 pond. Every two months, juvenile tilapia(45 g) were harvested and the pond was stocked withnew fingerlings (8 g). After six months, the total produc-tion was 63,000 tilapia juveniles and 282 kg ha¡1 ofprawns with mean weight of 2.4 g and a survival of80.4%.

The coculture of caged lambari (Astianax altipara-nae), a small Characidae fish within a pond stocked withM. rosenbergii at 5 PL m¡2 was also studied in Brazil bySussel et al. (2011). Lambari densities were 300, 450, and600 m¡3. After 60 days, the density of 300 fish m¡3

resulted in higher mean weight and length, but the bestproductivity was achieved at the highest density. As theobjective was the sale of lambari as live bait and the sur-vival did not differ significantly, the density of 600 m¡3

showed more profitability. Prawns weighed 3.8 g with asurvival of 91.4%, which would allow selling them asjuvenile IIs, with a productivity of 45,700 ha¡1. Recently,intensive research on the polyculture of M. amazonicumwith Nile tilapia and native fish in a semi-intensive sys-tem has been developed in Brazil. Data suggest that the

culture can be performed in fresh or low-salinity water(<2 salinity) and that the prawns do not affect tilapiadevelopment (Henry-Silva et al., 2015). Rodrigues (2013)evaluated the use of natural and artificial substrates inthe polyculture ofM. amazonicum with Nile tilapia. Thisauthor found that the use of substrates significantlyincreased mean prawn biomass by 34% and alsoincreased the number of prawns with individual massand length greater than 3.1 g and 7 cm, respectively.

Trials on fish-prawn polyculture have also been con-ducted in other countries. In an economic study per-formed in Panama, Engle (1987) reported higherinternal rates of return for two polyculture systems—Nile tilapia, grass carp, and M. rosenbergii; and a speciesof pacu (Colossoma mitrei; currently named Piaractusmesopotamicus), grass carp, and M. rosenbergii (17 and13%, respectively)—than in prawn monoculture (10%).In Saudi Arabia, Siddique et al. (1996) found that totalyield from the polyculture of Nile tilapia, common carpand M. rosenbergii was almost five times as great as thatof prawn monoculture. In a limited experiment (onepond; two cages) in Argentina, Wicki et al. (1998)showed that the coculture of pacu (Piaractus mesopota-micus) in cages in a pond stocked with M. rosenbergiiand grass carp seemed viable, resulting in fish survival of100%, fish production of 800 kg ha¡1, and nearly»780 kg ha¡1 of prawns per cycle (survival 89%).

In Puerto Rico, Garc�ıa-P�erez et al. (2000) comparedthe monoculture of 1.3 g prawns stocked at 7 m¡2 andthe monoculture of all-male 7.4 g Nile tilapia stocked at1 m¡2 with a polyculture system where 1.1 g prawns and7.4 g tilapia were stocked at 7 and 1 m¡2, respectively. Inan experiment lasting 145 days a commercial feed wasapplied at rates based on biomass (5% during days 1–90and 3% for days 91–120); maximum daily feeding ratesreached 50 kg ha¡1 in monoculture (prawns or fish) and67 kg ha¡1 in polyculture. Total yields and mean weightsof tilapia were not significantly different in polycultureor monoculture, but the yields of prawns were signifi-cantly different. Total prawn yield in monoculture was1367 kg ha¡1, but only 951 kg ha¡1 in polyculture. Finalprawn mean weights were 55 g in monoculture and 31 gin polyculture. The authors postulated that the fish mayhave efficiently consumed the major portion of the feedprovided and suggested that further studies should con-sider increasing feeding rates and improving feed distri-bution. These results also suggest that the wastesprovided by the tilapia were not sufficient to meet thefeed requirements of prawns stocked at 7 m¡2.

In Trinidad, Souza et al. (2005) obtained a maximummean production of 12,622 kg ha¡1 (prawns plus fish) inthe polyculture of M. rosenbergii with armored catfish(Hoplosternum littorale) and 24,176 kg ha¡1 in M.

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rosenbergii-tilapia polyculture, after 5.5 months of rear-ing. In the Philippines, Rola (2007) analyzed three differ-ent feeding systems (intensive, semi-intensive andtraditional) on the commercial production of M. rose-nbergii and milkfish (Chanos chanos) in polyculture, andfound better economic indicators for the semi-intensiveand intensive systems. In a trial conducted in Egypt,Sadek and Moreau (1996) studied the effect of stocking0.3 g PL or 2 g juvenile prawns at 2 and 5 m¡2 in mono-culture with the stocking of 0.3 g PL at 2 m¡2 in polycul-ture with two stocking densities of <1 g Nile tilapia (1 or2 m¡2) and 15 g common carp (0.25 or 0.5 m¡2). Thehighest annual rates of return were obtained from themonoculture of 2 g PL at 2 m¡2 (57.5%) and the polycul-ture of 0.3 g PL at 2 m¡2 with low-density tilapia andcarp (30.9%). Also in Egypt the effect of monocultureand polyculture of M. rosenbergii with Nile tilapia fry innursery tanks on the growth and survival of prawns wasexamined by El-Sheriff and Ali Mervat (2009). Theseauthors concluded that the polyculture system was mosteffective at a density of 100 prawns m¡2.

Rice-prawn culture

In an early study, Sadek and Moreau (1998) stocked M.rosenbergii at either 10,000 or 20,000 ha¡1 in monocul-ture and at 10,000 ha¡1 with 5000 ha¡1 of tilapia in poly-culture in rice paddies. Average prawn size at harvest didnot differ significantly between the two monoculturetreatments but was significantly reduced in polyculture.The maximum return (US$ 4507 ha¡1) was obtainedfrom the high-density prawn monoculture integratedwith rice culture; unsurprisingly, the lowest was obtainedby the rice-only culture treatment (US$ 904 ha¡1). Thisresearch has not been followed up and there is little orno freshwater prawn culture of any sort in Egypt at pres-ent (S. Sadek, personal communication, 2015).

In Thailand, Giap et al. (2005) compared different fer-tilization systems (basal and regular) and feeding regimes(with and without commercial feeding) in rice-prawnculture. The treatment with basal fertilization and com-mercial feed was more profitable, with higher yields ofrice (0.42 kg m¡1) and prawns (347 kg ha¡1; 23.8 g inindividual size) per crop.

Lan et al. (2006a) working in the Mekong Delta ofVietnam, concluded that in the rotational rice-prawnsystem, stocking densities of 1, 2, or 3 PL m¡2 producedgreater prawn mean weights, but densities of 3 or 4 PLm¡2 resulted in higher yields. The best economic viabil-ity was obtained at a density of 3 PL m¡2. There were nosignificant differences between the two feed manage-ments employed (commercial pellets and a combinationof pellets and snail meat) on production parameters;

however, the combination diet showed better net profitand cost-benefit ratio. In another experiment, Lan et al.(2006b) compared the two systems of culture (rotationaland integrated) and concluded that the rotational systemgave significantly higher profits than the integrated, butrequired a greater initial investment. Under both sys-tems, a stocking density of 2 PL m¡2 resulted in a signifi-cantly higher profit and cost-benefit ratio than 4 or 6 PLm¡2. In subsequent observations from farmer trials, Lanet al. (2008) noted that net profit was significantly higherfor a feeding management using pellets, trash fish, andsnail meat when compared to feeding with pellets only;in these trials the prawn stocking densities used were 4and 5 m¡2.

The polyculture of M. rosenbergii with fish in ricefields has been studied in depth in Bangladesh. Wahabet al. (2008) tested four densities of M. rosenbergii(10,000, 15,000, 20,000, and 25,000 ha¡1) in polyculturewith mola at a stocking density of 2 m¡2 in simultaneousrice-prawn-fish culture. Production ranged from 222–388 kg ha¡1 of prawns, 51–68 kg ha¡1 of mola and2880–3710 kg ha¡1 of rice. The prawn density of15,000 ha¡1 resulted in significantly higher productionof both prawn and mola, with a net profit of US$1100 ha¡1. In a similar experimentally designed study,Rohul Amin and Salauddin (2008) found that the inclu-sion of M. rosenbergii and mola in rice fields had pro-found impacts on the availability of nutrients in thewater and soil, increasing the yield of rice. The best har-vest of prawns (456 kg ha¡1) was achieved at a stockingdensity of 15,000 ha¡1 and the best production of rice(3710 hg ha¡1) was obtained at the prawn density of10,000 ha¡1. In another very similar study, Kunda et al.(2008) found that prawn production (mean survival 49–57%), which ranged from 294–596 kg/ha, was signifi-cantly higher in the treatment where 20,000 ha¡1 offreshwater prawns were stocked, compared with 10,000,15,000, or 25,000 ha¡1. This treatment also resulted inthe highest net profit margin (74%) indicating that20,000 ha¡1 prawns and 20,000 ha¡1 of mola is the bestcombination for prawn-mola culture in rain-fed ricefields after rice cultivation. Ali et al. (2009) did not findsignificant differences between the net profit fromprawn-carp polyculture and polyculture integrated withrice and vegetables growing on the banks (US$ 835 and857 ha¡1, respectively).

In India, Mishra and Mohanty (2004) studied differ-ent weir heights and densities of prawns and fish in rice-fish-prawn polyculture, concluding that short-durationfish and prawn rearing (about 120 days) with a totalstocking density (four species of fish with a 10% inclu-sion of prawns) of 25,000 ha¡1 and a weir height of12.5 cm resulted in the best net profit. Mohanty et al.

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(2004) evaluated the integration of prawns and fish inrice fields and concluded that the percentage increase inrice yield was 7.9–8.6% above rice monoculture, probablydue to the better aeration of water and additional supplyof fertilizer in the form of leftover feed and prawnexcreta. Mohanty (2010) studied the impact of periodiccull harvesting in rice-fish-prawn culture on the popula-tion structure, feed intake pattern and growth perfor-mance of M. rosenbergii, concluding that it enhances thenet return by 28%.

In Brazil, Marques et al. (2011), in a partial budgetanalysis, concluded that the introduction of M. rosenber-gii in rice-monoculture systems resulted in an increase of10.5% in total crop value. Boock et al. (2013) pointed outthat the native species M. amazonicum can be reared inrice fields without being fed and sold as live bait in thesport fishing market. These authors observed that thestocking of prawns in the rice fields did not degradeeffluent quality and sometimes improved it, probablybecause prawns consume organic matter, contributing tothe recycling of nutrients. Boock et al. (2016) concludedthat rice-prawn culture at a stocking density of 2 juve-niles m¡2 of M. rosenbergii without commercial dietswas economically feasible and could compensate for lowrice prices. Stocking at a higher rate (5 juveniles m¡2)was not cost effective due to an increase in prawn mor-tality and the higher cost of purchasing postlarvae. Theseauthors observed that natural food is not a major limit-ing factor for stocking densities of up to 5 prawns m¡2;prawns can attain commercial size in this system. In Bra-zil, the selling price of rice fluctuates and rice monocul-ture often becomes non-profitable. Thus, the integratedrice-prawn system may be an effective alternative for theproduction of rice without the necessity for governmentsubsidies.

Aquaponics and other systems

There are only two reports of studies on integrated sys-tems other than fish-prawn polyculture or coculture andrice-prawn culture. In Brazil, the effect of the use of efflu-ents from a M. amazonicum nursery, operated at a den-sity of 80 PL m¡2, in the production of hydroponicwatercress and lettuce was evaluated by Castellani et al.(2009). The results showed that the effluent was suffi-cient to meet the demand for nutrients only in the caseof watercress, when supplemented with a nutritive solu-tion. Further research is necessary to produce a moreconclusive result. In Bangladesh, Hoq et al. (1999)obtained good results from integrating poultry, prawnsand fish. These authors reported that the use of freshchicken manure from poultry sheds situated directlyover ponds stocked with M. rosenbergii and several fish

species (silver carp, grass carp, catla, rohu, mrigal, andThai silver barb Puntius gonionotus) resulted in signifi-cantly higher fish productivity (5290 kg ha¡1 in10 months) than the use of treated manure brought frombeyond the farm (3365 kg ha¡1 in 10 months) or thecontrol without fertilization (885 kg ha¡1 10 months).Prawn survival and production was significantly lower inthe ponds with fresh chicken manure droppings. Resultsshowed the potential to produce fish and prawns withoutfeed and to produce chicken above ponds, saving space.

Future potential for integrated freshwaterprawn farming in a sustainable world

Zimmermann et al. (2010) observed that the polycultureof fish is common in tropical aquaculture, but freshwaterprawn polyculture with fish is less common, despite theconsiderable amount of research on this subject. As oneof the possible reasons for this situation, New (2002)stated that the management of polyculture systems ismore complex than that of monoculture, especiallyregarding to the harvesting of prawns, since it is difficultto synchronize both fish and prawn production in orderto achieve the maximum production of marketable ani-mals. On the other hand, some of the constraints thatwere pointed out by Zimmermann et al. (2010) as limit-ing the expansion of integrated prawn cultures in bothAsia and Latin America, are the same as those that con-strain the increase of prawn monoculture in thoseregions, i.e., lack of technology transfer, shortages ofprawn PL and capital, and deficiencies in marketing.

In a review of the polyculture and coculture of fresh-water prawns with tilapia species, New and Valenti(2016) stated that both systems are technically feasibletoday and can improve the use of natural resources(such as space, water, and nutrients), reduce the use ofcommercial diets, and increase profits. Nevertheless, asignificant research effort is needed to provide science-based knowledge in order to improve the efficiency ofsuch aquaculture systems and to take full advantage oftheir potential.

Some of the difficulties experienced by producers andworkers in implementing fish-prawn polyculture couldbe minimized by using newly available technologies. Forexample, the undesirable proliferation of tilapia in prawnponds could be avoided by using sex-reversed or cagedtilapia. Problems in harvesting prawns and fish or thepolyculture of species with different culture cycles couldalso be solved by using caged fish, which can be har-vested independently from the harvest of prawns(Tidwell et al., 2000; Danaher et al., 2007).

The benefits that result from the implementation ofrice-prawn farming are already clear-cut. Ahmed and

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Garnett (2010) stated that the sustainability of rice-prawn farming in Bangladesh could be expressed interms of production technology (benefits of rice-prawnintegration that leads to an increase in rice production aswell as other diverse products such as prawns, fish, andvegetables), socioeconomic aspects (technology easilyassimilated by small-scale producers, increasing theirsocial status and food consumption—mainly proteina-ceous food—and creating thousands of direct and indi-rect jobs) and environmental aspects (although it causesan irredeemable loss of biodiversity, rice-prawn farmingreduces the use of pesticides and fertilizers, increases soilfertility and promotes other important environmentalbenefits). These authors added that rice-prawn farmingplays an important role in the economy of Bangladeshand that, although not entirely sustainable, it can helpthe country keep pace with the current demand for food.In addition Kamal (2010) considered that gher farminggenerates an average income that is four times higherthan any other typical agriculture practice in Bangladeshand increases the demand for labor, reduces food insecu-rity, provides education, protein, health care, andsanitation.

Conclusions

The global landscape of integrated freshwater prawn cul-ture can be considered as falling into three categories:

1. Having great social and economic importance, as isthe case of rice-prawn culture in some Asiancountries.

2. Being well studied and having established a theo-retical basis for implementation, but still needingto be commercially expanded, as is the case ofprawns and fish growing together in a pond(polyculture).

3. Having great promise but needing more studies tobe effectively adopted by the production sector,which is the farming of caged fish in prawn ponds(coculture).

Other integrated systems, such as prawn aquaponics,still depend on the development of viable technologies.The effective adoption of integrated prawn culture byproducers will depend much more on strengthening andorganizing the freshwater prawn production chain thanon technical solutions, in the same way as currentlyoccurs in freshwater prawn monoculture.

Tilapia and prawns are compatible species for farmingthrough either polyculture or coculture. Both systems aretechnically feasible today and can improve the use of nat-ural resources (such as space, water, and nutrients),reduce the use of commercial diets, and increase profits.Nevertheless, a significant research effort is needed to

provide science-based knowledge in order to improvethe efficiency of such aquaculture systems and to takefull advantage of their potential.

There are good prospects for the expansion of inte-grated freshwater prawn culture worldwide. Apart fromthe high market value of freshwater prawns and theincreasing interest in prawn production, especially inAsia and South America, some other factors that maylead to this expansion are as follows:

1. There is a worldwide trend by consumers in select-ing products cultured in a sustainable way. Inte-grated systems are recognized as being moreefficient in using natural resources, especiallyspace, feed, and water, thus being more sustainable.The dissemination of these qualities may be a goodmarketing strategy that leads to obtaining a pre-mium price for products from integrated systems.

2. The semi-intensive monoculture of tropical fish inponds often shows minimal economic viability,especially in small-scale or family-run systems. Onthe other hand, many of these species can be inte-grated with prawns, resulting in profitable systemswhen carried out efficiently, even on a small scale.

3. Caged tilapia culture in lakes and reservoirs isgradually replacing pond culture in some tropicalcountries, such as Brazil. Due to this, there aremany abandoned or underutilized natural ponds,which could be turned into profitable fish-prawnrearing units.

4. Integration of prawns into rice monoculture canincrease the profitability, mainly in small-scale sys-tems. Furthermore, the production of organic orfree-chemical rice is generally a small-scale enter-prise compatible with prawn integration and pre-mium prices are paid for rice produced in thesesystems.

5. The farming of fish confined in cages installedwithin ponds stocked with free-swimming prawns(coculture) is a promising system. This systemoffers some advantages when compared with therearing of fish and prawns together (polyculture).This includes the possibility of producing variousspecies of fish at the same time in the same prawnpond, which would allow production to be adaptedto address niche markets. Further research, includ-ing studies on the carrying capacity of the ponds,and economics is still essential.

Acknowledgments

The authors wish to express their sincere gratitude to CynthiaVilar Boock for providing artwork for our review.

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Funding

The authors are also grateful to FAPESP and CNPq for finan-cial support and for the fellowships granted.

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