Pen culture of Macrobrachium amazonicum : use of artificial diet and impact on benthic community

Pen culture of Macrobrachium amazonicum: use of

artificial diet and impact on benthic community

Fabr�ıcio Martins Dutra1, Yara Moretto2, Leandro Portz3 & Eduardo Luis Cupertino Ballester1,3

1Postgraduate Program in Zoology, Federal University of Paran�a, Curitiba, Paran�a, Brazil2Federal University of Paran�a - Sector Palotina, Palotina, Paran�a, Brazil3Postgraduate Program in Aquaculture and Sustainable Development, Federal University of Paran�a - Sector Palotina,

Palotina, Paran�a, Brazil

Correspondence: F M Dutra, Universidade Federal do Paran�a- Setor Palotina, Sala de P�os-graduac�~ao, Bloco Semin�ario, Rua

Pioneiro, 2153, Jardim Dallas, Palotina, Paran�a 85950-000, Brasil. E-mail: [email protected]

Abstract

The pressure on the benthic macroinvertebrate com-

munity and the use of artificial diets were evaluated

during the pen rearing of Macrobrachium amazoni-

cum. The study followed a completely randomized

design with three treatments and three repetitions:

treatment no prawns, no feed (noPnoF); treatment

prawns, no feed (PnoF); and treatment prawns, feed

(PF). Six pens were stocked with 10 prawns m�2

with average initial weight of 0.63 � 0.05 g and

average length of 4.28 � 0.14 cm; the other three

pens were not stocked with prawns. Every 10 days,

prawn biometry was carried out and samples were

collected from the water and the sediment in each

pen. The results showed that the limnological vari-

ables remained adequate for the species’ biology.

The zootechnical indices did not differ significantly

(P > 0.05) in weight, total length and survival, with

values of 4.43 � 0.93 g, 8:03 � 0:43 cm, and

80 � 9.6%, respectively, for the treatment PnoF,

and 4.83 � 1.03 g, 8.42 � 0.51 cm and 73 �3.5% for treatment PF. The macroinvertebrate

analyses showed a significant difference (P < 0.05)

for the total abundance of Oligochaeta among the

days of collection and for Polymitarcyidae among

the treatments. It is therefore concluded that the

presence of prawns affects the abundance in the

benthic community and that the zootechnical per-

formance of M. amazonicum in this experimental

condition is not influenced by supplied artificial feed.

Keywords: trophic relations, pen culture, ben-

thic invertebrates, Macrobrachium amazonicum,

fresh water prawn

Introduction

Macrobrachium amazonicum is a species with great

production potential and important zootechnical

characteristics such as quick growth and rusticity

(Moraes-Valenti & Valenti 2010). Moreover, it has

high nutritional and gastronomic value, being

greatly accepted in international markets

(Damasceno, Andrade & Stamford 2009). Thus,

M. amazonicum has great socioeconomic and envi-

ronmental importance in the north and northeast

regions of Brazil (Silva, Fr�edou & Filho 2007; Mor-

aes-Valenti, Moraes, Preto & Valenti 2010) and is

the freshwater prawn species most widely explored

by artisanal fishing (Maciel & Valenti 2009). How-

ever little is known about its nutritional require-

ments and feeding behaviour under culture

conditions.

Studies on the feeding behaviour of prawns

show that, even in a production system with arti-

ficial feed supply, natural feeding represents a sig-

nificant part of these animals’ diet. Corbin,

Fujimoto and Iwai (1983) suggest that most of

the macronutrients needed for freshwater prawns

must be supplied by prepared feeds, but the

required micronutrient levels may derive from

natural productivity in ponds. Anderson, Parker

and Lawrence (1987) estimated that between

53% and 77% of the growth of Litopenaues vanna-

mei came from grazing natural food items present

in ponds.

According to Shishenchian and Yusoff (1999),

descriptions of the stomach content indicated that

among the natural foods available, benthic fauna

is an important food source in prawn farming

© 2014 John Wiley & Sons Ltd 1

Aquaculture Research, 2014, 1–10 doi:10.1111/are.12488

ponds. Benthic macroinvertebrates comprise a

group of aquatic organisms with size starting at

1 mm (Day, Hall, Kemp & Y€a~nez-Arancibia 1989).

Most belong to the Arthopoda, Mollusca, Annelida,

Nematoda and Platyhelminthes phyla, involving a

wide range of species found in almost all types of

freshwater habitats under different environmental

conditions (Eaton 2003). The benthic community

is an important component in the sediment of riv-

ers and lakes and is crucial for the nutrient

dynamics and transformation of matter into

energy flow (Rosa, Freitas & Niencheski 2009). It

represents one of the main and essential food

sources of the aquatic fauna, directly contributing

to the ecology dynamics (Rosa et al. 2009).

According to Moss (2002), semi-controlled envi-

ronments may be used in order to clarify ecologic

and biological issues, since in this type of system

the energy input and output can be quantified and

there are no restrictions in sampling regarding the

time and space variations as is the case of natural

environments. Soares, Peixoto, Bemvenuti, Wasi-

elesky, D’inc~ao, Murcia and Suita (2004) state

that using pens as experimental units in natural

water bodies or in ponds also enables investigating

aspects related to the animals feeding ecology and

zootechnical performance.

Therefore, the present study aimed at assessing

the pressure imposed on the benthic macroinverte-

brate community deriving from pen farming of

M. amazonicum and the contribution of artificial

feed on prawn performance.

Materials and methods

This study was carried out in a pond in a commer-

cial farm in the city of Palotina, PR, Brazil (24°160

53.97″ S, 53°500 24.61″ W). The post-larvae

M. amazonicum used in the experiment were pro-

vided by the Freshwater Prawn Farming Labora-

tory of the Center of Aquaculture of CAUNESP, of

the S~ao Paulo State University Julio de Mesquita

Filho, in Jaboticabal, SP, Brazil.

Experimental design

Nine round pens were used with 5 mm mesh and

10 m2 of bottom area fixed in the bottom of the

pond for a total area of 0.6 ha and 1.47 m of

average depth (Fig. 1). The experiment lasted for

60 days and the M. amazonicum juveniles with

average weight of 0.63 � 0.17 g and average

length of 4.28 � 0.34 cm were stocked in six

randomly chosen pens at a density of 10

prawns m�2. The study used a completely ran-

domized design with three treatments and three

repetitions: treatment prawn, no feed (PnoF); treat-

ment prawn, feed (PF), and no prawn, no feed

(noPnoF). In each pen, three initial sediment sam-

ples were collected for the analysis of abundance

and composition of the benthic community and

granulometric analysis.

Over the experimental period, the animals in the

PF treatment received supplemental feeding twice

a day (11:00 and 16:00 hours) with commercial

pelleted prawn feed (Guabi� Potimar), with 35%

crude protein at a feeding rate equivalent to 10%

of the total biomass (Marques, Barros, Mallasen,

Boock & Valenti 2012; Kimpara, Moraes-Valenti,

Queiroz & New 2013). Every 10 days, prawn

biometry, water collection and sediment collection

were performed for quantitative and qualitative

macroinvertebrate analysis and granulometry

analysis.

Prawn biometry

Every 10 days, 10 prawns were collected from each

treatment with the aid of a dip net and the wet

weight (0.01 g) and total length (digital caliper)

were determined for each animal. After biometry,

the prawns were taken back to their treatments.

Benthic macroinvertebrates

The sampling of the initial benthic macroinverte-

brate community was performed on the first day

of the experiment, before animal stocking (D0) and

before prawn biometric sampling (D10; D20; D30;

D40; D50; D60). Three sediment samples were col-

lected from each treatment with a Petersen collec-

tor with area of 0.042 m2 (Davanso & Henry

2006).

The benthic community was pre-sorted in a sys-

tem of granulometric sieves with different mesh

sizes (2.0 mm, 1.0 mm, 0.5 mm, 0.25 mm). After

pre-sorting, they were transported to the Labora-

tory of Ecology, Fishing and Ichthyology of the

Federal University of Paran�a – Campus Palotina

(LEPI-UFPR) for sorting, counting and stereoscopic

microscope identification. The animals were pre-

served in 70% alcohol and were identified accord-

ing to the taxonomic keys by Mugnai, Nessimian

and Baptista (2010).

© 2014 John Wiley & Sons Ltd, Aquaculture Research, 1–102

Pen culture of M. amazonicum F M Dutra et al. Aquaculture Research, 2014, 1–10

The total benthic macroinvertebrate fauna was

estimated using the following formula:

Individuals=m2 S � AAc

Individuals/m2, Average amount of benthos per

m2; S, Average amount of collected benthos; A,

Total area of cultivation; Ac, Total area of the

collector.

Sediment analysis

Every 10 days, sediment samples were collected

from each pen with a Petersen collector in order

to determine the organic matter content and sedi-

ment granulometry.

After drying at ambient temperature, the sam-

ples were weighed and taken to a furnace at 600°C for 3 h and weighed again to determine the

organic matter content of the sediment (Suguio

1973). The granulometry analysis was carried out

according to Wentworth (1922).

Water quality monitoring

The water quality variables were measured every

10 days inside the pens at average depth of 1 m

for dissolved oxygen (Hanna® HI 9146 oxygen

meter, Woonsocket, RI, USA), temperature (Incon-

term digital thermometer), and pH (PHTEK 100

pH meter), while water transparency was deter-

mined with a Secchi disk.

Water samples were stored in 1 L polyethylene

bottles in a refrigerated environment and sent to

the Laboratory of Water Nutrition, Bromatology

and Quality for Aquatic Organisms of the Federal

University of Paran�a – Campus Palotina (UFPR),

where the concentrations were determined for

nitrite according to Mackereth, Heron and Talling

(1978) and ammonia according to Koroleff

(1976), besides alkalinity and hardness according

to Walker (1978).

Statistical analyses

For the statistical analysis, the data were sub-

jected to normality verification through the

Shapiro–Wilk test and to homogenicity verification

through Levene’s test. When these two require-

ments were met, ANOVA was applied followed by

Tukey’s test (P < 0.05), while when they were

not met, mathematical transformations were

performed. A two-way ANOVA (treatment 9 time)

was carried out to identify significant variations in

the abundance of benthic organisms (logx+1), in

sediment granulometry, and percentage of organic

matter using the software Statistica 7.0 (Statsoft

Inc. 2004).

The statistical analysis of the zootechnical indi-

ces was carried out only in the treatments PnoF

and PF, for which Student’s t-test (P < 0.05) was

used for weight (g), length (cm), and final sur-

vival (%).

Results

Prawn growth monitoring

Over the experimental period, temperature ranged

from 26.67 to 31.29°C. Dissolved oxygen had

higher values in the treatment PF (6.04 �1.80 mg L�1) than in the treatments noPnoF

(5.85 � 2.02 mg L�1), and PnoF (5.92 �1.80 mg L�1). The pH values fluctuated slightly

and remained close to neutral. The other variables

had similar values among the treatments

(Table 1).

Table 1 Mean and standard deviation (�SD) of limno-

logical variables in treatment no prawn, no feed

(noPnoF), treatment prawns, no feed (PnoF) and treat-

ment prawn, feed (PF) for 60 days

Variables

Treatments

noPnoF PnoF PF

Temperature

H2O

28.90 � 2.18 28.90 � 2.23 29.00 � 2.29

Depth (cm) 1.47 � 0.02 1.47 � 0.00 1.47 � 0.01

Transparency

(cm)

28.95 � .3.47 28.95 � .3.45 28.95 � .3.46

pH 7.60 � 0.76 7.60 � 0.75 7.60 � 0.72

Dissolved

Oxygen

(mg L�1)

5.85 � 2.02 5.92 � 1.80 6.04 � 1.80

Alkalinity

(meq L�1)

25.31 � 13.65 25.31 � 13.65 25.31 � 13.65

Hardness

(mg L�1)

22.39 � 9.51 22.39 � 9.51 22.39 � 9.51

Nitrite

(mg L�1)

0.021 � 0.016 0.021 � 0.016 0.021 � 0.016

Ammonia

(mg L�1)

0.062 � 0.103 0.062 � 0.103 0.062 � 0.103

*Mean values (�SD) did not differ significantly (P > 0.05)

among treatments.

© 2014 John Wiley & Sons Ltd, Aquaculture Research, 1–10 3

Aquaculture Research, 2014, 1–10 Pen culture of M. amazonicum F M Dutra et al.

The final survival did not significantly differ

(P > 0.05) between treatments (Table 2). The

average weight and total length also did not differ

significantly (P > 0.05) at the end of the experi-

ment, with weight 4.43 � 0.93 g for the treat-

ment PnoF and 4.83 � 1.03 g for the treatment

PF. The total length was around 8.03 � 0.43 cm

in treatment PnoF and 8.42 � 0.51 cm in

treatment PF (Fig. 1).

Benthic macroinvertebrate community monitoring

365 222 individuals were recorded overall for the

benthic macrofauna studied (Table 3). The analy-

sis of its composition revealed the presence of five

main orders (Rhynchobdellida, Mesogastropoda,

Ephemeroptera, Odonata and Diptera) and one

class (Oligochaeta). Within these orders, eight fam-

ilies were identified (Glossiphoniidae, Ceratopogoni-

dae, Chaoboridae, Chironomidae, Polymitarcyidae,

Gomphidae, Libellulidae and Ampullariidae).

Oligochaeta was the taxon occurring the most

and with the greatest total abundance (86.69%)

among the treatments, while for the other taxa

total abundance was less notable, as seen for Chi-

ronomidae (10.48%), Polymitarcyidae (1.89%),

Glossiphoniidae (0.40%), Ceratopogonidae (0.33%)

and Chaoboridae (0.14%). Ampullariidae was

found only in noPnoF and PnoF, responding for

0.05% of the total abundance. Gomphidae and Li-

bellulidae were restricted to noPnoF (0.04%;

Table 3).

No significant difference (P > 0.05) was found

for the average total abundance values of ben-

thic macroinvertebrates among the treatments

over the experimental period. The only exception

found was for PF after 10 days of the beginning

of the experiment (D10), when a greater varia-

tion was found in benthic organism abundance

(Fig. 2).

The families Ampullariidae, Chaoboridae, Gom-

phidae and Libellulidae represented 0.23% of the

total abundance. Thus, due to their low abun-

dance, these families were not considered for the

statistical analysis. All other families were counted

and analysed separately among the treatments.

However, the comparison among the total abun-

dance of each taxon was verified individually

among the treatments and over time.

The class Oligochaeta was found in all treat-

ments (Table 3) and was the most abundant,

reaching densities above 5000 individuals/m2 at

D40. The abundance for this class did not show a

significant difference (P > 0.05) among the treat-

ments over time, but had a significant difference

(P < 0.05) among the treatments within each time

period, being representative in the treatments at

D40 and D60 (Fig. 3).

The abundance of the family Glossiphoniidae

was not significantly influenced (P > 0.05) among

the treatments over time. However, the test of var-

iance could not be performed to detect significant

differences among the treatments over time due to

the absence of this family in certain periods of the

experiment (Fig. 4).

Table 2 Mean and standard deviation for IBW, FBW, IBL, FBL, survival, FC and total biomass of Macrobrachium ama-

zonicum in the treatments prawn, no feed (PnoF) and treatment prawn, feed (PF) for 60 days

Trataments IBW (g) FBW (g) IBL (cm) FBL (cm) Survival (%) FC (g) Biomass total (g)

PnoF 0.63 � 0.05* 4.43 � 0.93 4.28 � 0.14 8.03 � 0.43 80 � 9.6 – 352.97 � 48.79

PF 0.63 � 0.05 4.83 � 1.03 4.28 � 0.14 8.42 � 0.51 73 � 3.5 1.5:1 354.20 � 16.96

*Mean values (�SD) did not differ significantly (P > 0.05) between treatments.

IBW, initial body weight; FBW, final body weight; IBL, initial body length; FBL, final body length; FC, feed conversion.

2010 60504030Days

00

1

2

3

4

5

6

7

8

9

10

Bod

y w

eigh

t (g)

Bod

y le

ngth

(cm

)

PnoF PF

PnoF PF

Figure 1 Mean body weight and mean body length

(�SD) of Macrobrachium amazonicum reared in pen sys-

tem in two different treatments (Treatment prawn, no

feed – PnoF and Treatment prawn, feed – PF) during

60 day of the experiment.



The family Ceratopogonidae did not show a

statistical difference (P > 0.05) for abundance

over time. On the other hand, the significance

(P < 0.05) could not be verified among the treat-

ments at each time due to the absence of indi-

viduals from this family in the samples of

certain experimental periods. This absence is well

noted in the treatments PnoF, regardless of feed,

from the experimental period D30 until D60

(Fig. 5).

The family Chironomidae was found in all treat-

ments (Table 3) and was the second most

abundant family in number of individuals/m2.

However, the abundance in Chironomidae did not

have a significant difference (P > 0.05) among the

treatments over time and within each period

(Fig. 6).

For the family Polymitarcyidae, significant differ-

ences (P < 0.05) were found for the treatments

PnoF and PF over time. However, the significance

could not be tested for the treatments within each

–10 0 10 20 30 40 50 60Days

0.0

0.5

1.0

1.5

2.0

2.5

3.0

3.5

4.0

4.5

5.0

Indi

vidu

als

m–2

(log

x+1 )

noPnoF PnoF PF

a a

b

Figure 2 Mean abundance (individuals/m2 = logx+1)

of benthic macroinvertebrates during pen rearing for

the treatment no prawn, no feed (noPnoF), treatment

prawns, no feed (PnoF) and treatment prawn, feed

(PF).

0

10 000

20 000

30 000

40 000

50 000

60 000

70 000

80 000

90 000

0 10 20 30 40 50 60

Indi

vidu

als

m–2

Days

Oligochaeta

noPnoF

PnoF

PFb

a a

a

bb

Figure 3 Abundance (individuals/m2) of the Oligocha-

eta class over the experimental period (days) among

the treatment no prawn, no feed (noPnoF), treatment

prawns, no feed (PnoF) and treatment prawn, feed

(PF).

Table 3 Composition and total abundance of benthic macroinvertebrate community (n) and their frequency of occur-

rence (%) in the treatment no prawn, no feed (noPnoF), treatment prawns, no feed (PnoF) and treatment prawn, feed

(PF)

Taxa

Occurrence of benthic community among treatment

noPnoF, n (%) PnoF, n (%) PF, n (%) Total, n (%)

Annelida

Rhynchobdellida

Glossiphoniidae 667 (0.18) 389 (0.11) 389 (0.11) 1444 (0.40)

Oligochaeta* 61 833 (16.93) 112 889 (30.91) 141 889 (38.85) 316 611 (86.96)

Arthropoda

Diptera

Ceratopogonidae 667 (0.18) 333 (0.09) 222 (0.06) 1222 (0.33)

Chaoboridae 333 (0.09) 111 (0.03) 56 (0.02) 500 (0.14)

Chironomidae 15 778 (4.32) 13 389 (3.67) 9111 (2.49) 38 278 (10.48)

Ephemeroptera

Polymitarcyidae 6611 (1.81) 167 (0.05) 111 (0.03) 6889 (1.89)

Odonata

Gomphidae 56 (0.02) – – 56 (0.02)

Libellulidae 56 (0.02) – – 56 (0.02)

Mollusca

Mesogastropoda

Ampullariidae 111 (0.03) 56 (0.02) – 167 (0.05)

Total 86 111 (23.58) 127 333 (34.86) 151 778 (41.56) 365 222 (100)

*Rating to Class.



period due to the absence of individuals in certain

experimental periods. On the other hand, it can be

seen that the abundance of this family was greater

in noPnoF (Fig. 7).

Granulometric composition

The granulometric fraction of the soil had a preva-

lence of sandy sediment. Throughout the experi-

mental period, the coarse sand and organic matter

were the only fractions that did not significantly

differ (P > 0.05). Among the treatments, no signif-

icant differences (P > 0.05) were found for med-

ium sand, mud, or organic matter, while the other

fractions had significant differences among the

treatments (P < 0.05; Table 4).

Discussion

Limnologic variables

The limnologic variables remained within the

appropriate range for the biology of M. amazoni-

cum. According to Sampaio, Silva, Santos and

Sales (2007), this species is found in water with

temperature between 27.5 and 31°C, with dis-

solved oxygen from 4.6 to 6.1 mg L�1, and pH

between 7.4 and 8.4. Timmons, Ebeling, Weathon,

Summerfelts and Vinci (2002) recommended total

ammonia levels below 3 mg L�1 and nitrite below

1 mg L�1 in warm-water aquiculture. New (2002)

recommended a range of 20–60 mg L�1 of CaCO3

for M. rosenbergii and transparency between 25

and 60 cm.

Prawn growth monitoring

Prawn growth did not differ significantly between

the treatments for weight and length, which was

contrary to the initial hypothesis of a greater gain

in weight and length in the treatment PF. Hence,

0

20

40

60

80

100

120

140

160

180

200

0 10 20 30 40 50 60

Indi

vidu

als

m–2

Days

Glossiphoniidae noPnoF

PnoF

PF

Figure 4 Abundance (individuals/m2) of the family

Glossiphoniidae over the experimental period (days)

among the treatment no prawn, no feed (noPnoF),

treatment prawns, no feed (PnoF) and treatment

prawn, feed (PF).

0

50

100

150

200

250

300

350

0 10 20 30 40 50 60

Indi

vidu

als

m–2

Days

Ceratopogonidae

noPnoF

PnoF

PF


Ceratopogonidae over the experimental period (days)



prawn, feed (PF).

0

1000

2000

3000

4000

5000

6000

7000

8000

0 10 20 30 40 50 60Days

Chironomidae

noPnoF

PnoF

PF

Indi

vidu

als

m–2


Chironomidae over the experimental period (days)



prawn, feed (PF).

0

500

1000

1500

2000

2500

3000

0 10 20 30 40 50 60

Days

Polymitacyidae

noPnoF

PnoF

PF

Indi

vidu

als

m–2


Polymirtacyidae over the experimental period (days)



prawn, feed (PF).



it is supposed that the natural food is able to

maintain prawn growth for several weeks with lit-

tle or no input of supplementary feeding. Correia,

Pereira, Apolin�ario, Horowitz and Horowitz (2002)

noted a significant increase in M. rosembergii

growth in ponds with natural feeding. Lanari, Bal-

lestrazzi and Tibaldi (1989) found favourable

growth for Marsupenaeus japonicus reared in ponds

with low stocking rate (<10 m�2) regardless of the

input of allochthonous feeding.

Prawn survival in the present study was similar

to the one suggested by Valenti, New, Salin and

Ye (2010), who reported 50–80% survival for

M. rosenbergii in semi-intensive ponds using a

stocking rate of 4–12 prawn m�2. Similar values

to those in the present study were reported by

Souza, Stringuetta, Bordingon, Bohnenberger, Bos-

colo and Feiden (2009) in a polyculture study

with M. amazonicum and Oreochromis niloticus.

Preto, Pizzato and Vallenti (2008), on the other

hand, found that the same species reared during

100 days with artificial feeding had average sur-

vival of 85.5% at the same stoking density (10

prawn m�2). Marques et al. (2012) found survival

Table 4 Granulometric composition (%) of soil among the treatment no prawn, no feed (noPnoF), treatment prawns,

no feed (PnoF) and treatment prawn, feed (PF): evaluating the fraction of pebbles (4.00 mm), granules (2.00 mm), very

coarse sand (1.00 mm), coarse sand (0.500 mm), medium sand (0.250 mm), fine sand (0.125 mm), very fine sand

(0.063 mm), mud (<0.063 mm) and organic matter (OM) over the experimental period

Granulometry/

treatment

Days

D0 D10 D20 D30 D40 D50 D60

4.00 mm

noPnoF 0 � 0 0 � 0 0 � 0 0 � 0 0 � 0 0.07 � 0.12 0 � 0

PnoF 0 � 0 0 � 0 0 � 0 0 � 0 0.21 � 0.36 0 � 0 0 � 0

PF 0 � 0 0 � 0 0 � 0 0 � 0 0 � 0 0.50 � 0.75 0 � 0

2.00 mm

noPnoF 3.17 � 3.31 6.07 � 6.68 6.64 � 0.81 2.38 � 0.51 063 � 0.68 0.28 � 0.11 0.89 � 0.61

PnoF 2.41 � 1.31 2.33 � 1.91 6.23 � 0.42 2.48 � 2.03 0.96 � 0.60 0.07 � 0.10 0.92 � 0.63

PF 2.34 � 1.60 1.30 � 0.30 2.73 � 2.43 2.48 � 1.21 0.76 � 0.52 1.70 � 1.43 1.39 � 0.42

1.00 mm

noPnoF 15.74 � 9.36 14.07 � 2.73 14.29 � 1.30 10.18 � 0.99 7.41 � 2.83 5.98 � 0.99 7.97 � 3.52

PnoF 11.16 � 3.05 12.68 � 4.50 12.91 � 1.74 10.74 � 2.87 5.67 � 1.77 3.76 � 0.72 7.86 � 2.76

PF 13.48 � 4.10 12.41 � 1.11 6.63 � 4.88 10.54 � 1.40 5.92 � 2.36 9.54 � 4.16 8.71 � 2.72

0.500 mm

noPnoF 13.06 � 2.89 9.86 � 2.69 10.99 � 0.36 14.15 � 2.73 13.36 � 3.69 12.50 � 1.39 11.97 � 3.37

PnoF 10.68 � 2.86 9.80 � 0.59 10.13 � 1.35 12.18 � 2.50 10.63 � 3.16 9.26 � 2.38 11.65 � 1.52

PF 11.18 � 2.24 9.12 � 1.00 8.04 � 2.79 12.58 � 1.50 11.71 � 2.89 14.44 � 4.58 11.81 � 2.08

0.250 mm

noPnoF 9.93 � 0.57 7.44 � 1.72 9.03 � 0.64 7.53 � 5.03 11.76 � 1.44 11.53 � 0.54 10.51 � 2.31

PnoF 10.32 � 3.7 8.90 � 2.80 8.36 � 0.83 10.63 � 2.51 10.56 � 3.10 9.72 � 2.86 10.54 � 0.93

PF 9.24 � 1.82 7.97 � 1.37 9.31 � 2.05 10.96 � 1.30 13.15 � 1.35 13.99 � 3.65 10.99 � 1.46

0.125 mm

noPnoF 11.11 � 2.91 9.29 � 1.42 10.67 � 1.29 10.48 � 0.53 11.20 � 0.94 11.58 � 0.61 11.14 � 1.75

PnoF 12.61 � 3.74 12.42 � 3.99 10.05 � 0.97 11.85 � 2.92 13.73 � 0.75 12.31 � 2.11 11.62 � 0.98

PF 10.64 � 2.02 11.59 � 3.31 12.30 � 1.76 12.41 � 1.55 15.96 � 4.45 14.55 � 2.00 13.25 � 2.72

0.063 mm

noPnoF 17.62 � 0.77 25.94 � 4.21 15.82 � 1.17 14.64 � 1.51 15.51 � 2.11 17.62 � 2.50 17.31 � 3.59

PnoF 18.67 � 3.40 25.94 � 9.51 16.65 � 0.92 15.14 � 3.29 20.81 � 4.37 21.08 � 4.43 16.72 � 2.81

PF 17.38 � 3.79 30.59 � 4.22 19.74 � 3.81 15.97 � 0.89 18.21 � 2.53 15.73 � 4.79 17.35 � 3.74

<0.063 mm

noPnoF 18.50 � 12.19 14.45 � 8.42 17.41 � 2.28 22.57 � 1.80 22.73 � 3.00 24.83 � 1.43 26.42 � 8.48

PnoF 21.53 � 9.37 16.72 � 2.54 19.97 � 1.61 23.12 � 4.69 24.29 � 5.00 29.49 � 5.39 27.05 � 3.55

PF 21.59 � 12.00 14.84 � 2.81 27.44 � 8.82 22.12 � 2.67 22.00 � 4.24 18.56 � 8.45 23.67 � 4.75

OM

noPnoF 10.35 � 3.56 16.13 � 0.45 12.92 � 1.84 13.23 � 0.19 16.17 � 4.61 13.51 � 2.56 12.65 � 0.81

PnoF 11.83 � 2.91 10.77 � 0.98 15.56 � 3.30 13.01 � 2.16 11.43 � 1.60 11.99 � 2.20 12.13 � 1.37

PF 12.99 � 1.77 10.74 � 0.91 12.24 � 3.86 11.75 � 1.01 10.95 � 1.89 9.52 � 2.61 11.86 � 1.61

Data represent mean values (�SD).



above 90% when studying the influence of stock-

ing rate (20 prawn m�2) in the nursery phase on

the growth of M. amazonicum reared in pens. The

high survival rates found in the study may be due

to the low stocking rate and lack of predators.

By the end of the experiment, females in repro-

ductive process (ovigerous) were found in the

treatment PF. The same was not found in the

treatment PnoF, which indicates that, even

though natural feeding supplies the needs of

prawns over certain periods, artificial feeding still

carries essential elements to prawn development.

Benthic community monitoring

According to the results obtained, a clear reduc-

tion in the abundance of some macroinvertebrate

species was found over the experimental period

and among the treatments. That indicates that the

prawns are able to adapt, increasing predation on

the benthic community when there is no exoge-

nous feeding.

Oligochaeta and Chironomidae were the most

abundant and most occurring taxa since they

were present in all treatments and throughout the

experimental period. This result was also found by

Tidwell, Webster, Sedlacek, Weston, Knight, Hill,

D’abramo, Daniels, Fuller and Monta~nez (1955) in

a study with complete and supplementary diets

and organic fertilization in the production of

M. rosenbergii, when the researchers found a prev-

alence of Oligochaeta. Santos, Terra, Junior, S�a

and Junior (2011) state that the high abundance

of these classes is associated to the high organic

matter levels. Rodrigues (2003), in a study in a

reservoir in the medium and low portions of the

Tiete river, concluded that Oligochaeta tend to

increase in abundance compared to Chironomidae,

while Bemvenuti (1987) stated that the inverte-

brate macrofauna strategically bury themselves

and/or maintains high densities as mechanisms to

evade predation.

Callisto, Moretti and Goulart (2001) considered

that the presence of Oligochaeta and Chironomidae,

which were found in the present study, indicate

poor water quality. However, Polymitarcyidae,

which is a highly sensitive family that does not tol-

erate high trophic status, were also recorded in the

present study. Leal and Esteves (2000) associate the

presence of Polymitarcyidae to lentic environments

with clear waters and the presence of fine particles

in the sediment of the aquatic ecosystems, as was

the case of the experimental conditions in the pres-

ent study.

Polymitarcyidae and Ceratopogonidae were

more abundant mainly in the treatment noPnoF,

which may indicate predation processes or sedi-

ment disturbance caused by grazing. Karr (1999)

reported that processes of severe disturbance in

aquatic systems lead to the loss of benthic life

sustainability and the consequent complete elimi-

nation of aquatic organisms in these environ-

ments.

Other studies showed that benthic organisms

decrease in abundance over the rearing period likely

due to the pressure imposed by the prawns (Shi-

shenchian & Yusoff 1999; Soares et al. 2004). Ti-

dwell et al. (1995) claim that prawns increase

predation when there is no exogenous feeding.

Moreover, the experimental handling of fish popula-

tions has shown that biomass and production rates

of benthic macrofauna drastically decreased with

the increase in predation pressure (Kajak 1970).

Sediment composition and benthic fauna

Significant differences (P < 0.05) were found in

the sediment fraction during the experimental per-

iod among treatments. Clark, Frid and Attrill

(1997) stated that the association of benthic macr-

oinvertebrates can be influenced by physical char-

acteristics of the different types of substrata found

in aquatic environments.

According to Uieda and Gajardo (1996), sedi-

ment composition can be an important factor for

the maintenance of the benthic community since

it provides food and shelter. Soares et al. (2004)

mentioned that the predatory action of prawns on

benthic invertebrates may also disturb the substra-

tum by revolving it.

Allan and Castillo (2007) described that, even

in substrata with particles smaller than 1 mm,

small invertebrates thrive and serve as food. So-

ares et al. (2004) mentioned that the organic mat-

ter in rearing is normal and results from the

decomposition of organic matter from the exoge-

nous feeding, metabolic processes and prawn

waste. Therefore, the exogenous feeding may have

influenced the variation in the percentage grain

size over time and among treatments. However,

one cannot claim that this variation has influ-

enced the benthic community since the presence

of prawns imposes pressure on the macroinverte-

brate benthic community.



Conclusion

The results of this study indicated that the zoo-

technical performance of M. amazonicum reared at

densities of 10 prawns m�2 in areas with high

availability of benthic macrofauna can be main-

tained with no added feed for 60 days.

The study also confirmed that the presence of

prawns reduces the abundance of Ceratopogoni-

dae, Chironomidae and Polymirtacyidae over time

in the treatments stocked with prawns.

Acknowledgments

The authors thank Mr Carlos Piovesan for provid-

ing the experimental pond and the Laboratory of

Freshwater Prawn Aquaculture Center of the Uni-

versidade Estadual Paulista J�ulio de Mesquita Filho

(CAUNESP) for the donation of M. amazonicum

post-larvae. Professor Eduardo L.C Ballester. and

Professor Leandro Portz are research fellows of

CNPq Agency. This work was developed with

CNPq and FINEP funding.

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Pen culture of Macrobrachium amazonicum : use of artificial diet and impact on benthic community

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