Feeding rhythms and diet of Farfantepenaeus paulensis under pen culture in Patos Lagoon estuary, Brazil

www.elsevier.com/locate/jembe

Journal of Experimental Marine Biolog

Feeding rhythms and diet of Farfantepenaeus paulensis under pen

culture in Patos Lagoon estuary, Brazil

R. Soaresa,b,T, S. Peixotob, W. Wasieleskyb, F. D’Incaoa

aLaboratorio de Crustaceos Decapodos, Fundacao Universidade Federal do Rio Grande (FURG), Departamento de Oceanografia,

C.P. 474, Rio Grande (RS), 96201-900, BrasilbLaboratorio de Maricultura, Fundacao Universidade Federal do Rio Grande (FURG), Departamento de Oceanografia, C.P. 474,

Rio Grande (RS), 96201-900, Brasil

Received 16 November 2004; received in revised form 23 February 2005; accepted 23 February 2005

Abstract

The feeding habits of Farfantepenaeus paulensis under pen culture in the Patos Lagoon estuary (Brazil) were studied. A

total of 1074 shrimp had their stomach contents analyzed. Samples were taken bi-hourly over a 24 h period at about 10 day

intervals during 2 months. F. paulensis did not exhibit strict feeding periodicity (except at D21–22 and D32–33). Thus, the offering

of feed pellets during daylight is recommended but the rates of feeding must be further investigated. F. paulensis showed an

omnivorous feeding behavior. Despite daily supply of feed pellets, natural food comprised the major part of the shrimp stomach

contents. Among prey organisms, polychaetes and tanaids were the main groups recorded. Consumption of detritus and plant

material decreased as shrimp grew. Intake of feed pellets increased significantly in the second month of culture (i.e. 5 g mean

shrimp weight), thus it might be possible to reduce the initial input of commercial feed.

D 2005 Elsevier B.V. All rights reserved.

Keywords: Farfantepenaeus paulensis; Feeding behavior; Patos Lagoon; Pen shrimp culture; Stomach content analysis

1. Introduction

The pink shrimp Farfantepenaeus paulensis

(Perez-Farfante) is one of the most valuable species

captured in Patos Lagoon estuary (Southern Brazil),

0022-0981/$ - see front matter D 2005 Elsevier B.V. All rights reserved.

doi:10.1016/j.jembe.2005.02.019

T Corresponding author. Laboratorio de Maricultura, Depto. de

Oceanografia-FURG, C.P. 474, Rio Grande (RS), CEP: 96201-900,

Brasil. Fax: +55 53 2336601.

E-mail address: [email protected] (R. Soares).

but fisheries production has been decreasing in recent

years mostly due to over-fishing of the estuarine and

oceanic populations (Reis and D’Incao, 2000; D’In-

cao and Reis, 2002; D’Incao et al., 2002). Therefore,

considering the potential of the estuarine areas for

aquaculture development, pen culture of F. paulensis

in estuarine shallow waters has been proposed as an

additional income for artisanal fishermen residing in

the shore areas (Wasielesky et al., 2003).

In aquaculture systems, feeding times must be

adjusted to coincide with shrimp activity periods to

y and Ecology 322 (2005) 167–176

R. Soares et al. / J. Exp. Mar. Biol. Ecol. 322 (2005) 167–176168

reduce the leaching of nutrients and feed accumulation

(Cuzon et al., 1982). Most penaeids spend the day

buried in the sediment and emerge and feed at night

(Dall et al., 1990). Usually the circadian light rhythm

controls food intake of decapods when they are not

subjected to tidal cycles, however additional diurnal

intake can occur to complement nocturnal feeding

according to metabolic requirements (Reymond and

Langardere, 1990). Experiments under semi-intensive

culture conditions revealed different and variable

feeding behavior among species. For example, Far-

fantepenaeus subtilis showed day and night feeding

activity (Nunes et al., 1996); Penaeus monodon shifted

feeding activity pattern from night to day-time during

culture (Focken et al., 1998), while Marsupenaeus

japonicus shifted from continuous to nocturnal feed-

ing as shrimp grew (Reymond and Langardere, 1990).

Wild F. paulensis juveniles captured in the Patos

Lagoon estuary showed a more intensive feeding

activity during the dark period (Santos, 2003); how-

ever there is no previous investigation of F. paulensis

feeding behavior under pen culture.

Penaeids have been described as opportunistic

omnivores (Dall et al., 1990). In earthen ponds,

natural food can comprise a significant part of the

diet and nutrition of shrimp even when pelleted food

is provided (Hunter et al., 1987; Reymond and

Langardere, 1990; Allan et al., 1995; Nunes et al.,

1997; Focken et al., 1998; Nunes and Parsons, 1999).

Anderson et al. (1987) estimated that between 53%

and 77% of Litopenaeus vannamei growth was due to

the grazing on pond biota. Reports from shrimp gut

contents have demonstrated a range of dietary items in

addition to artificial food in culture ponds, such as

polychaetes, amphipods, nematodes, copepods,

bivalves, diatoms and detritus (Allan and Maguire,

1992; Allan et al., 1995; Nunes et al., 1997; Focken et

al., 1998; Martinez-Cordova et al., 1998). During the

pen culture of F. paulensis in the Patos Lagoon

estuary (summer months), the shrimp have access to a

variety of natural food items, including detritus, plant

material and animal prey (Soares et al., 2004).

Artificial feed is also provided during the pen culture,

but the contribution of natural and commercial feed to

the diet of F. paulensis is not known. Therefore, the

present study aimed to investigate the feeding rhythms

and the diet of F. paulensis reared in pen enclosures in

Patos Lagoon estuary.

2. Materials and methods

2.1. Experimental pens and shrimp feeding

This study was carried out in a shallow estuarine

area in the Patos Lagoon, Rio Grande do Sul State,

Brazil (32803V55n S, 52812V30n W) over 64 days.

Experimental juvenile shrimp (0.90F0.02 g body

weight) were obtained from spawnings of wild

broodstock conducted at the Marine Aquaculture

Center (Estacao Marinha de Aquacultura-EMA, Uni-

versity of Rio Grande).

Three 50 m2 pens (8 m diameter and 2 m height)

made with polyester net covered by PVC (5 mm

mesh) were set up in the estuary and used as

experimental units. Pens were randomly stocked

with 26 juvenile/m2. Shrimp were fed twice daily

(0900 hours and 2100 hours) with commercial

pelleted feed (35% crude protein level) (Camaronina

Purina, Sao Lourenco da Mata, PE, Brazil). Initial

feeding rate was 15% of shrimp biomass/day, but

from day 18 to the end of the study, this rate was

reduced to 5%.

Shrimp growth was evaluated on D10 (i.e. 10 days

after shrimp stocking), D21, D32, D42, D52 and D63,

sampling 50 shrimp from each pen, which were

weighed and returned to the pens. Feeding ration was

adjusted based on average shrimp body weight.

Temperature and salinity were monitored daily during

the culture period.

2.2. Shrimp sampling and feeding rhythms

Shrimp were sampled for stomach content anal-

ysis at D10, D21, D32, D42, D52 and D63 using a

push-net. Five shrimp from each pen were captured

bi-hourly over 24 h periods. Animals were fixed and

preserved in 70% ethanol solution for further

analysis. Sampling period extended from 1000 hours

to 0800 hours on the following day. A total of 180

animals per sampling day were collected (i.e. 5

shrimp�3 pens�12 sampling times). Soft shrimp

were not used for analyses and were returned to the

pens.

In the laboratory, shrimp were sexed and measured

(carapace length). Animals were then dissected and

their proventriculus was removed. The degree of

fullness was determined visually by positioning the

R. Soares et al. / J. Exp. Mar. Biol. Ecol. 322 (2005) 167–176 169

proventriculus over a light box. The stomach repletion

index (rs) was estimated as a percentage of fullness as

follows (adapted from Reymond and Langardere,

1990):

Table 1

Water quality parameters and shrimp weights from each sampling

day (D10–11 to D63–64) during the estuarine pen culture of

Farfantepenaeus paulensis

Sampling

days

Temperature

(8C)Salinity

(x)

Mean (FSD)

body weight (g)

D10–11 28 5 1.7F0.04

D21–22 27 5 3.3F0.07

D32–33 22 4 4.1F0.08

D42–43 21 8 5.1F0.10

D52–53 22 7 5.3F0.10

D63–64 17 7 5.4F0.09

Degree of

fullness

0 (empty) nearly 0 N0�H

full

NH�O

full

NO�J

full

full

rs (%) 0 5 25 50 75 100

The stomach repletion rate (Rs) was calculated as:

Rs=PNrs/N, where rs=stomach repletion index for

each shrimp and N =total number of stomach ana-

lyzed during the period. Similarly, stomach repletion

rates for diurnal and nocturnal samples (Rs day and Rs

night) were computed according to day-time or night-

time period. All samples collected from 8:00 h to

18:00 h were considered as day times (n =6). Like-

wise, catches from 2000 h to 0600 h were considered

as night times (n =6).

Empty stomachs index ( fv) and full stomachs index

( fr) were also determined: fv (%)=100 V/N, where

V=number of empty stomachs; fr (%)=100 R/N

where R =number of full stomachs.

Comparisons between day and night stomach

repletion at the different sampling periods were

assessed by two-tailed t-tests.

2.3. Food items analysis

Food items analyses were performed on stomachs

of F. paulensis individuals collected at two day

times (1200 hours and 1400 hours) and two night

times (2400 hours and 0200 hours). The proven-

triculus was cut and contents were carefully washed

with distilled water into a petri dish. The contents

were left immersed in distilled water for a few

minutes to allow hydration and partitioning of the

items. Following this, contents were displaced over

a counting chamber gradated in 1 mm squares

(Sedgewick Rafter Cell) and examined through a

stereoscopic microscope. The abundance of each

food item was estimated by the area occupied in the

counting chamber (mm2). Stomach contents were

classified into 5 categories: prey (whole or frag-

mented animal body parts); plant material (seeds,

algae and aquatic macrophytes); detritus (fine

organic particles); pellets (brownish compact mate-

rial) and minerals (silt and sand grains). Prey items

were taxonomically classified, prey frequency (Cn)

and prey occurrence index ( f) were calculated as

follows:

Cn(%)=100� ( p/P), where p is the total amount

of each specific type of prey ingested (mm2) and P

is the total amount of prey ingested (mm2).

f(%)=100� (Np/N), where Np is the number of

stomachs with a specific prey and N is the total

number of non-empty stomachs.

3. Results

3.1. Water quality and shrimp growth

The daily mean temperature was 23 8C (F3.4)

and ranged from 17 to 29 8C. Higher values were

observed in the initial phase, but it decreased

progressively during the experiment. Salinity

remained low during the entire rearing cycle and

ranged from 3 to 9x (meanFSD=6.0F1.5). Mean

values of water quality parameters (i.e. temperature,

salinity) for each sampling day are presented in

Table 1.

Shrimp growth was low and after 64 days of

culture they attained 5.4 g (mean body weight) (Table

1). From D42–43 until the end of the rearing period

shrimp growth was negligible.

3.2. Feeding rhythms

A total of 1074 stomachs were analyzed. No strict

feeding pattern was observed and stomachs with food

were recorded from all sample times (Fig. 1).

D63- D64

0

20

40

60

80

100

10 12 14 16 18 20 22 24 2 4 6 8

Time (h)

D21- D22

0

20

40

60

80

100

D32- D33

0

20

40

60

80

100

10 12 14 16 18 20 22 24 2 4 6 8

Time (h)

D42- D43

0

20

40

60

80

100

D52- D53

0

20

40

60

80

100

Sto

mac

h r

eple

tio

n (

%)

D10 - D11

0

20

40

60

80

100

Fig. 1. Variation (meanFSE) of stomach repletion rate (Rs) of Farfantepenaeus paulensis during day and night (shaded area) sampling periods

(D10–11 to D63–64).


On D10–11, feeding activity was intense over the

entire 24 h sampling cycle (Fig. 1). The mean stomach

repletion rate (Rs=77.8%) was the highest value

p = 0.026N=195

p = 0.000*N=186

p = 0.480N=178

0

20

40

60

80

100

D10-11 D21-22 D32-33

Sampl

Sto

mac

h r

eple

tio

n r

ate

(%)

Rs day Rs

Fig. 2. Mean (+SE) diurnal (Rs day), nocturnal (Rs night) and overall (Rs) sto

period (D10–11 to D63–64). Values above bars indicate results from t-tes

samples.

recorded during the experiment. The stomach fullness

was similar ( p N0.05) between day (Rs day=76.4 %)

and night times (Rs night=79.4%) (Fig. 2). These

p = 0.794N=167p = 0.598

N=167

p = 0.085N=180

*

D42-43 D52-53 D63-64

ing days

night Rs

mach repletion rates of Farfantepenaeus paulensis for each sampling

ts. Asterisks denote significant differences between day and night

0

10

20

30

40

50

D10-11 D21-22 D32-33 D42-43 D52-53 D63-64

Sampling days

Em

pty

an

d f

ull

sto

mac

h in

dex

(%

)

fv fr

Fig. 3. Empty stomachs index ( fv) and full stomachs index ( fr) of

Farfantepenaeus paulensis in the sampling periods (D10–11 to

D63–64).


values indicate that food consumption was

continuous throughout the light and dark periods.

The highest value of full stomachs index

(fr =47.8%) and the lowest empty stomachs index

Fig. 4. Volumetric representation of food items and vacuity in Farfantepe

sampling periods (D10–11 to D63–64). Empty stomachs were not included.

(fv =6.7) confirm the intense feeding activity

(Fig. 3).

On D21-22 and D32-33 feeding activity was lower

than on D10-11 especially during the daytime, indicat-

ing higher feeding intensity during the dark period

(Fig. 1). Consequently, Rs night became significantly

higher ( p b0.05) than Rs day (Fig. 2). The fr dropped

and fv rose on D21–22. Similar values were observed

on D32–33 (Fig. 3).

On D42–43, feeding activity increased in both night

and day periods (Fig. 1). The Rs increased to 66.4%

and Rs night was higher than Rs day but with no

significant difference ( p N0.05) (Fig. 2). The increase

on feeding activity reduced the fv and strongly

improved fr values (Fig. 3).

On D52–53 and D63–64, feeding activity was lower

than on D42–43 and the differences between night and

day food consumption became even less evident

(Figs. 1 and 2). The values of fr were lower than on

D42–43 but fv remained constant (Fig. 3).

naeus paulensis stomach contents in day and night samples in the

Table 2

Occurrence index f (%) and frequency of prey items Cn (%) in the stomach contents of Farfantepenaeus paulensis cultured in estuarine pens for

each sampling period (D10–11 to D63–64)

Sampling period D10–11 D21–22 D32–33 D42–43 D52–53 D63–64

Prey f Cn f Cn f Cn f Cn f Cn f Cn

Polychaeta 89.7 38.4 100.0 38.0 100.0 31.9 86.7 40.9 88.9 59.6 100.0 71.4

Crustacea

Tanaidacea 86.2 25.7 79.3 27.0 73.9 27.7 83.3 35.8 74.1 17.5 75.0 12.9

Ostracoda 86.2 12.8 72.4 9.7 69.6 16.1 70.0 6.4 51.9 3.7 54.2 5.1

Gastropoda

Heleobia sp. 69.0 11.1 58.6 7.7 43.5 8.7 20.0 9.2 33.3 12.2 33.3 8.5

Prey fragments 65.5 11.9 82.8 17.6 52.2 15.5 46.7 7.6 33.3 7.0 25.0 2.0


3.3. Food items analysis

Stomach contents from 360 shrimp sampled over

the rearing period were analyzed. The diet consisted

of varied food items, but natural food (i.e. plant

material, detritus and animal prey) was more abundant

than commercial food (feed pellets) at all sampling

times (Fig. 4).

3.3.1. Detritus

Detritus was the major food component in

the diet of F. paulensis and represented more

than 35% of the stomach contents during the

rearing period. The abundance of detritus in the

0

50

100

Pre

y o

ccu

rren

ce (

%)

Polychaeta Tanaidacea Ostra

D10-11 D21-22 D32-33

Sampl

Pre

y fr

equ

ency

(%

)

0

100

50

Fig. 5. Occurrence ( f) and frequency (Cn) indexes (%) of prey items

paulensis cultured in pen enclosures. PF=prey fragments. Empty stomach

stomach contents decreased slightly as shrimp grew

(Fig. 4).

3.3.2. Plant material

Plant material was consumed, but in small amounts.

The highest values were observed at D10–11 especially

from day samples, but the ingestion of plant material

tended to reduce as shrimp grew (Fig. 4).

3.3.3. Prey

Animal prey was an important food item in the diet

of F. paulensis. In general, prey consumption fluc-

tuated along the culture period and tended to decrease

as shrimp increased in size (Fig. 4). Pronounced

coda Gastropoda PF

D42-43 D52-53 D63-64

ing days

by taxonomic groups in the stomach contents of Farfantepenaeus

s were not included.


differences in prey abundance in stomach contents

were observed between day and night samples, which

were found to be related to the rhythm of feeding

activity. For example, on D21–22 the abundance of

prey in the stomachs was 15% higher in the night

samples, indicating that F. paulensis applied a higher

predation pressure on the benthic fauna during peaks

of feeding activity.

The occurrence (f) and frequency index (Cn) of

prey items, categorized by taxon, are presented in

Table 2 and Fig. 5. Very small and unidentified animal

parts were classified as bprey fragmentsQ (PF).

Polychaetes and tanaids were the most important prey

items in all culture stages. Polychaetes were present in

100% of the stomachs analyzed on D21–22, D32–33 and

D63–64. Their frequency (Cn) rose along time and at

D63–64 more than 70% of the prey items were

polychaetes. Tanaids also had high occurrence and

represented an important fraction of the stomach

contents. Occurrence and frequency of the smaller

sized prey (i.e. ostracods, prey fragments and gastro-

pods) declined as the shrimp grew.

3.3.4. Minerals

Minerals comprised a minor fraction in the

stomach contents and the highest occurrences were

recorded from periods when shrimp feeding levels

were highest (i.e. D10–11 and D42–43) (Fig. 4).

3.3.5. Artificial food

Artificial food was an important portion of the

shrimp diet. Its abundance in the stomach contents

increased throughout the trial and exceeded the animal

prey ingestion in some occasions (Fig. 4).

Overall, analysis of the stomach contents indicated

that as shrimp grow the abundance of plant material

and detritus diminishes while the abundance of pellets

increases.

4. Discussion

4.1. Feeding rhythms

Variations on F. paulensis feeding behavior were

observed during the experimental rearing cycle. At the

beginning (D10–11), young juveniles showed intense

and continuous feeding regardless of light or dark

hours. After this period (D21–22 and D32–33), the Rs

was reduced and a significantly lower ingestion of

food was observed during daytime. In the second

month (D42–43), feeding activity became progressively

more even between day and night times until the end

of the culture period. Changes in the feeding behavior

during shrimp growth have been also documented for

other penaeid species. Both M. japonicus (Reymond

and Langardere, 1990) and Penaeus semisulcatus

(Heales et al., 1996) changed from continuous to

nocturnal feeding behavior as they grew. In contrast, P.

monodon shifted from night to day feeding activity

(Focken et al., 1998).

Daytime feeding activity was observed for wild-

caught Fenneropenaeus merguiensis (Chong and

Sasekumar, 1981), Litopenaeus setiferus (McTigue

and Feller, 1989), Penaeus esculentus (O’Brien,

1994), P. semisulcatus (Heales et al., 1996) and for

pond-reared F. subtilis (Nunes et al., 1996). Wild

juveniles of F. paulensis in the Patos Lagoon estuary

also appeared to feed over the entire 24 h cycle, but

feeding intensity increased significantly during the

dark period (Santos, 2003). However, F. paulensis

exhibited no strict feeding periodicity in the present

work, as shrimp were found to have similar day and

night indexes of stomach fullness for most of the

sampling days. This less pronounced periodicity may

be due to the offering of feed pellets in the morning,

which could be stimulating the feeding activity.

Similarly, peaks of food intake (both natural food

and pellets) in pond-reared F. subtilis occurred soon

after the spread of feed pellets (Nunes et al., 1996).

The constant presence of food in the stomachs of F.

paulensis may be in part attributed to the differential

digestibility of the diverse food items. Hyslop (1980)

pointed out that differential digestion rates for hard

and soft tissues may bias the results of stomach

contents analysis as the identification process relies

largely upon hard parts. Observations on digestion

rates of shrimp fed on soft tissues (i.e. crustacean and

bivalve meat) showed that P. monodon digested 53%

of its gut contents (Marte, 1980) and P. esculentus

emptied its stomachs within 1 h (Hill and Wassenberg,

1987). In a laboratory study, F. paulensis juveniles

(0.87–8.42 g) fed commercial feed pellets were able to

empty their stomachs after 3–4 h when deprived of

food (Soares et al., in press). This rapid rate of food

passage through the stomach allows F. paulensis to


feed continually or several times over a 24 h cycle and

the bias from differential digestion is probably small.

Thus, the offering of feed pellets during light period is

recommended for F. paulensis under estuarine pen

culture, but the rates of feeding must be further

investigated.

4.2. Shrimp diet

Penaeids have been described as opportunistic

omnivores (Dall et al., 1990) and this great opportun-

ism precludes shrimp from being placed in one trophic

group throughout their life cycle (Moriarty, 1976).

Likewise, in the present study F. paulensis showed an

omnivorous feeding behavior as it fed on several food

items throughout the culture period.

Minerals comprised only a small part (less than

4%) of the stomach contents. Its ingestion may be

involuntary and related to the consumption of food

items containing attached minerals. Small amounts of

minerals were also observed in the diet of P. monodon

(6.12%) (Marte, 1980), M. japonicus (less than 4%)

(Reymond and Langardere, 1990) and F. subtilis

(6.24-9.08%) (Nunes et al., 1997).

Detritus in penaeids stomachs appears to be almost

exclusively derived from plants (e.g. salt marshes and

seagrass beds), which is nutritionally enriched (e.g.

proteins and lipids) by the associated microbial food

web (Moriarty, 1976, 1997; Dall et al., 1990). Large

amounts of detritus are frequently observed in the

shrimp diet, but for some species the ingestion of

detritus could be just a supplement when other

preferred food items are scarce (Marte, 1980, 1982;

Chong and Sasekumar, 1981; Schwamborn and

Criales, 2000). In the present study, detritus was the

main food component of F. paulensis diet during the

entire rearing cycle. Likewise, in pond culture detritus

was the main diet constituent for P. monodon

(Bombeo-Tuburan et al., 1993) and comprised a

significant part of the diet of F. subtilis, especially

in younger juveniles (Nunes et al., 1997).

Although a high incidence of detritus in the stomach

contents of F. paulensis was observed in the present

study, the actual dietary importance of this food item

could not be determined. The refractory nature of

detritus may lead to its overestimation in stomach

contents, while more easily digestible organisms that

are consumed along with detritus may provide the bulk

of nutrition (Stoner and Zimmerman, 1988; Dittel et al.,

1997). The nutritional role of detritus is difficult to

assess but stable isotopes studies indicated that detritus

associated organisms were the main nutritional sources

for penaeids in estuarine areas (Stoner and Zimmer-

man, 1988; Sullivan and Moncreiff, 1990; Newell et

al., 1995; Dittel et al., 1997).

The amount of plant material consumed by

penaeids varies widely even within the same species

(Dall et al., 1990). The small amounts of plant material

found in F. paulensis stomachs are comparable to the

values recorded for F. subtilis (11%) (Nunes et al.,

1997) and Farfantepenaeus duorarum (7%)

(Schwamborn and Criales, 2000). In penaeids there

is a clear trend for juveniles to ingest more vegetable

matter than adults (Dall et al., 1990; Nunes et al., 1997;

Schwamborn and Criales, 2000). This pattern was also

observed in the present work for F. paulensis, as the

ingestion of plant material decreased from 8.4 (D10–11)

to 1.2% (D63–64) during daytime. The decreased

consumption of plant material and detritus throughout

the culture period suggests that F. paulensis juveniles

improve their predatory behavior as they grow.

During the rearing period F. paulensis juveniles

were observed to feed on distinct organisms, but

polychaetes and tanaids were the most significant prey

items in their diet. These two groups were also des-

cribed as the most abundant benthic invertebrates in the

pen culture of F. paulensis in the Patos Lagoon estuary

(Soares et al., 2004). In accordance, the diets of several

penaeids were coincident with seasonal and/or spatial

trends in the abundance of the major prey organisms,

which has been related to the shrimp’s opportunistic

behavior exerting a high predation pressure on a wide

range of available fauna (Chong and Sasekumar, 1981;

Stoner and Zimmerman, 1988). Soares et al. (2004)

demonstrated that the abundance of benthic macro

invertebrates dropped 86% after 21 days inside pen

enclosures where F. paulensis was cultured. Similarly,

results from the present study suggest that the reduction

on the total amount of prey consumed by F. paulensis

throughout the culture period was probably related to

the decline of prey availability in the pens.

The abundance of small prey items, i.e. ostracods,

gastropods and prey fragments, in the F. paulensis diet

reduced as shrimp grew. Likewise, in several penaeids

the nutritional importance of small sized food items

diminishes with ontogeny (Chong and Sasekumar,


1981; Stoner and Zimmerman, 1988; O’Brien, 1994;

Heales et al., 1996). According to Stoner and

Zimmerman (1988), ontogenetic shifts in the abun-

dance of prey organisms in the diet of penaeids are

undoubtedly related to increasing size of the chelae

and mouth parts. Therefore, the ingestion of small

sized prey organisms may decrease during F. paulen-

sis growth as their ability to capture and handle larger

and faster prey improves.

The importance of artificial food in the diet of F.

paulensis increased progressively throughout the

culture period. This may reflect an increasing require-

ment of shrimp for animal protein and the reduction of

prey’s availability to meet their nutritional needs.

Records on the consumption of natural food versus

artificial pellets in shrimp culture are variable as it

depends on several factors such as natural food

availability and feeding methods. For M. japonicus,

pelleted food intake did not exceed 4% of average

stomach volume (Reymond and Langardere, 1990),

whereas for F. subtilis over half of the diet was derived

from artificial food (Nunes and Parsons, 1999).

Artificial feeds may represent more than 50% of

the production costs in semi-intensive systems (Jory,

1995), thus the management of natural productivity

has been suggested to reduce feed inputs. Several

studies point out to the importance of pond fertiliza-

tion to stimulate the production of natural food to

shrimp nutrition (Lanari et al., 1989; Allan et al.,

1995; Jory et al., 2001; Martinez-Cordova et al.,

2003). Although this procedure is unviable in open

systems, such as estuarine pens, several organisms

occur naturally and may be exploited. Our results

indicate that it might be possible to reduce the initial

input of commercial diet, as a significant intake of

feed pellets was not observed until after shrimp

reached 5 g (i.e. second month of culture).

Acknowledgments

We would like to thank the staff from E.M.A. who

worked in the shrimp production, pen construction,

sampling and experiment maintenance. Thanks to Dr.

Ronaldo Cavalli for his revision and comments on this

manuscript. This study was financed by FAPERGS

(Fundacao de Amparo a Pesquisa do Estado do Rio

Grande do Sul, RS, Brazil). The first author was

supported by a doctorate scholarship from CNPq of the

Ministry for Science and Technology of Brazil. [RH]

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