Aquaculture Research, 2005, 36, 1075^1084

doi:10.1111/j.1365-2109.2005.01323.x

Biotic communities and feeding habits of Litopenaeus

vannamei (Boone 1931) and Litopenaeus stylirostris

(Stimpson 1974) in monoculture and polyculture
semi-intensive ponds
Luis R Mart| nez-Cordova1 & Emilio Pen
a-Messina2
1

Dictus, Universidad de Sonora, Hermosillo, Sonora, Mexico

Universidad Autonoma de Nayarit,Tepic, Nayarit, Mexico

Correspondence: L R Martinez-Cordova, Dictus, Universidad de Sonora, PO Box 1819, Hermosillo, Sonora 83000, Mexico. E-mail:
lmtz@guaymas.uson.mx

Abstract
Zooplankton, macrozoobenthos and feeding habits of
Litopenaeus stylirostris and L. vannamei in monoculture and polyculture semi-intensive experimental
ponds were evaluated. Zooplankton was more abundant in monoculture of L. stylirostris (1002  670
organisms (org.) L 1) than in monoculture of L.
vannamei (470  37 org. L 1), and polyculture
(321  188 org. L 1). The main zooplanktonic groups
were polychaeta larvae, nauplii, copepods and polychaeta. Macrozoobenthos was more abundant in
polyculture (6898  11137 org. m 2) compared with
monoculture of L. stylirostris (3201  350 org. m 2)
and L. vannamei (2384  3752 org. m 2). The main
benthic groups were copepods, polychaeta, ostracods,
nematodes and insects. Dierences in feeding habits
were found between species and regimes. Litopenaeus
vannamei showed to be a more voracious species and
fed mostly on organic detritus and benthos in both
culture regimes. Litopenaeus stylirostris had a more
restricted sources of feed in the ponds. The major component in the stomach content of both species was
detritus. Macroalgae, sand, exuvia, formulated feed,
prey and microalgae were minor components for both
species (o7%). Ingestion of formulated feed was
o4% for L. stylirostris and was not detected for L. vannamei. The stomach repletion rates were larger for L.
vannamei (55.6% and 48.8%) than for L. stylirostris
(43.75% and 44.89%). Litopenaeus stylirostris grew
better in polyculture (10.3  3.4 g) that in monocul-

r 2005 Blackwell Publishing Ltd

ture (9.0  3.8 g). Litopenaeus vannamei grew better

in monoculture (16.1  4.8 g) than in polyculture
(13.4  4.5 g). For both species, feed conversion ratio
was lower in polyculture.

Keywords: shrimp feeding, Litopenaeus vannamei,

Litopenaeus stylirostris, polyculture

Introduction
The Pacic white shrimp Litopenaeus vannamei
(Boone 1931) and the blue shrimp L. stylirostris
(Stimpson 1974) are the most commercially important Penaeid species in Mexico and the most popular
in American aquaculture. Nutrition of farmed
shrimp in semi-intensive systems is provided by formulated feed and the biota in the ponds (Mart| nezCordoba, Pasten-Miranda & Barraza-Guardado
1998). The reliance on these two nutritional sources
is a common practice in the culture of L. stylirostris
and L. vannamei alone or together (Lawrence 1985;
Goxe, Galinie & Ottogalli 1988; Villalon 1991). Formulated feed plays an important role in semi-intensive
production of shrimp (Rubright, Harrel, Holcomb &
Parker 1981; Garson, Pretto & Rouse 1986; Anderson,
Parker & Lawrence 1987; Hernandez-Llamas,
Hernandez-Lizardi, Gonzalez-Garibay & MagallonBarajas 1993; Hernandez-Llamas, MagallonBarajas, Lechuga-Deveze, Bustillos-Guzman & LopezCortes 1995; Martinez-Cordova, Campan
a-Torres &
1075

Feeding habits of white and blue shrimp L R Mart| nez-Cordova & E Pen
a-Messina Aquaculture Research, 2005, 36, 1075^1084

Porchas-Cornejo 2003). However, formulated feed

can represent 30^40% of production costs (UshaRani, Redly & Ravindranath 1993; Mart| nez-Cordova
1994; Jory 1995; Zendejas-Hernandez 2004).
Natural productivity in ponds, enhanced by fertilization, can help to minimize the use of supplemental
feed (Mart| nez Cordova 2000; Mart| nez-Cordova &
Campan
a-Torres 2000). The importance of biota in
the nutrition of farmed shrimp is well documented
(Marte 1980; Moriarty & Barclay 1981; Rubright et al.
1981; Hopkins, Blair, Grados, Maier, Sander &
Strokes 1988; Jory 1995; Martinez-Cordova, Campan
a-Torres & Porchas-Cornejo 2002). Natural food
may contribute more than 50% of the required nutrients for penaeid farmed shrimp (Anderson et al.1987;
Castille & Lawrence1989). Some studies have demonstrated low intake of formulated feed for some species
such as Marsupenaeus japonicus (Bate 1888) (Rollet
1986; Cam 1987; Reymond & Lagerdere 1990) and its
preference for organisms such as chironomid insects
(Gunderman & Poper 1977; Lumare, Scovacricchi,
Piscitelli & Grasso 1985). Other authors suggest that
the benthic nature of shrimp denes them as omnivorous, feeding on small benthic organisms and
organic detritus (Rubright et al. 1981). Lanari, Ballestrazzi and Tibaldi (1989) found a direct eect of pond
fertilization and stocking density on the feeding habits of M. japonicus, showing that this species adapts
its diet to food availability.
Litopenaeus vannamei and L. stylirostris exhibit
some dierences in their feeding preferences. Litopenaeus stylirostris is probably more carnivorous than
L. vannamei, which can consume plant material quite
well. These dierences suggest that a polyculture of
both species may result in the optimization of feed
sources in ponds.
Our objective was to analyse and describe the composition of zooplankton and benthos as well as the
gut contents of white shrimp L. vannamei and blue
shrimp L. stylirostris in semi-intensive monoculture
and polyculture regimes. This information may
contribute to optimizing feeding practices for these
species.

Materials and methods

The study was conducted over 10 weeks at the facilities of the Unidad Experimental Kino of the University of Sonora, northwest Mexico. A single-factor
experimental design with two replicates was used
for the study. Experimental units consisted of six

1076

earthen ponds (0.04 ha each) aerated from the bottom with a blower and a system of PVC pipes. Daily
water exchange was between 5% and 7%. All the
ponds were similarly managed. Three treatments
were evaluated: monoculture of L. vannamei
(MONO-V), monoculture of L. stylirostris (MONO-S)
and the polyculture of both species (POLYCUL-SV).
The initial stocking density was 30 postlarvae m 2 in all the treatments. In the polyculture
treatment, there were15 postlarvae m 2 of each species. Postlarvae were obtained from a hatchery in Bahia Kino, Sonora, Mexico located approximately
20 km from our facilities. Feeding in each pond was
adjusted weekly as a function of the growth of the organisms. A 25% crude protein-formulated feed was
used (Camaronina 25 using PurinaTM, Cargill Animal Nutrition, Ciudad Obregon, Sonora, Mexico). Previous studies have demonstrated that this protein
level is adequate for L. vannamei and sucient for L.
stylirostris if an adequate fertilization regime is applied (Martinez-Cordova et al. 2003). The daily ration
was 6% of the total biomass for the rst 2 weeks and
4% from the third week until the end of the culture.
For pond fertilization, urea (50 kg ha 1) and
triple super-phosphate or ammonium phosphate
(20 kg ha 1) were used. The frequency of fertilization
was established as a function of water transparency
in the ponds, taking a Secchi reading greater than
35 cm as a critical point.
Zooplankton and macrozoobenthos abundance
and composition in the ponds were evaluated weekly.
For zooplankton, a constant volume of 80 L of water
from each pond was ltered using a plastic cube provided with a window with a 64 mm sieve. An aliquot
of the sample was microscopically analysed in a
Sedgwick^Rafter counting chamber (PhycoTech, St
Joseph, MI, USA).
For macrozoobenthos analysis, pond sediment
samples were taken with a10.2 cm diameter PVC core
sampler introduced to a depth of approximately 5 cm
at sampling points selected at random. Macrozoobenthos samples were washed and ltered in sieves
of 0.5, 5 and 10 mm, and were then xed in 5% buffered formalin with Bengal Rose as the colourant for
counting and identication in the laboratory (Barraza-Guardado 1996; Mart| nez-Cordoba et al.1998). The
identication of zooplankton and macrozoobenthos
groups was made using the identication keys of
Newell and Newell (1963) and Brusca (1973).
The production variables evaluated during the experiment were mean weight, weight gain, survival,
yield and feed conversion ratio (FCR). Mean weight

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Aquaculture Research, 2005, 36, 1075^1084 Feeding habits of white and blue shrimp L R Mart| nez-Cordova & E Pen
a-Messina

and weight gain were recorded weekly in the dierent treatments by weighing 30 organisms from each
pond. Survival, yield and FCR were calculated at the
end of the study when shrimp were harvested,
counted and weighed.
The main variables of water quality recorded during the experiment were salinity, pH, dissolved
oxygen and temperature, using a portable Refractometer, a Pin-PointTM pH meter (American Marine,
Ridgeeld, CT, USA), and aYSITM digital oxygen meter
with integrated thermometer (YSI, Yellow Springs,
OH, USA). Gut contents of shrimp were analysed
weekly in each of the treatments. A sample of eight
shrimp from each monoculture pond and eight
shrimp of each species from each polyculture pond
were taken for this purpose.
Stomach content analysis began by determining
the minimum sample size. This was dened by plotting the number of components in the stomach
contents against the number of stomach samples.
Evaluation of the components was carried out by
means of microscopic analysis of the stomachs after
separation of the anterior and posterior chambers
from the rest of the digestive tract of the shrimp. Subjective methods proposed by Lagardere (1972) and
Poxton, Eleftheriou and MacIntyre (1983) were used
for evaluation of the stomach repletion index (rs). Indices of full stomachs (fr), empty stomachs (fv) and
global stomach repletion rate (Rsg) were estimated
according to Todd (1907), Arnaud and Hureau (1966)
and Hureau (1970).
For stomach-content analysis, shrimp in phase A-1
of the intermoult cycle were excluded (Drach &
Tchernigovtze 1967) because during this phase
they do not feed. The data obtained from each of the
variables were analysed for normality and homogeneity of variances to decide the type of statistical analysis to be used. For homogeneous and normal data,
the one-way analysis of variance was used, and for
data that did not conform to normal distribution and
the homogeneity of variances, the analysis of var-

iance by ranks of Kruskal^Wallis was used. A posteriori means test was conducted for each analysis using
a 95% signicance level. Statistical analysis was carried out using Statistica for Windows and Excel for
Windows.
Results
Zooplankton abundance in the dierent treatments
was variable over time. A greater total abundance
was found in the MONO-S treatment compared with
the other two treatments (Table 1). The most abundant groups of zooplankton were polychaeta larvae,
nauplii, adult copepods, juvenile polychaeta and
others (organisms whose appearance was irregular
during the sampling, and numerically insignicant
as compared with the above-named groups). This pattern was observed in the three treatments of the experiment (Table 2).
The total abundance of macrozoobenthos was
greater in the treatment POLYCUL-SV than in both
the monoculture treatments (Table 1). The most
abundant benthic groups were ostracods, nematodes, polychaeta, copepods and insects (Table 3).
The global stomach repletion rate (Rsg%), empty stomach frequency (fv%) and full stomach frequency
(fr%) are included in Table 4. Greater food consumption was measured for L. vannamei in both culture regimes, with values close to100% of stomach repletion
recorded from some weekly samples and a mean of
55.6% in monoculture and 48.8% in polyculture. For
L. stylirostris, the mean stomach repletion rates were
43.8% and 44.9% for mono and polyculture respectively. The highest full-stomach frequency indicies
(fr%) were observed for L. vannamei, reaching up to
50% in both regimes, whereas the maximum for L.
stylirostris was 37.5%. This last species had the highest empty stomach frequency index (fv%), with a
maximum of 50% in both regimes. This shows that,
in general, L. stylirostris consumed less food than L.
vannamei during the whole experiment.

Table 1 Comparison of zooplankton and macrozoobenthos abundances among treatments

Zooplankton (org. L
Zoobenthos (org. m

)
)

MONO-S

MONO-V

POLYCUL-SV

1002.16  670.73c
3200.80  350.34b

470.15  371.61b
2383.64  3752a

320.75  187.88a
6898.06  11 136.88c

Mean values  SD (Kruskal^Wallis 95%).

Dierent letters in a row indicates signicant dierences (Po0.05).
MONO-S, monoculture of Litopenaeus stylirostris; MONO-V, monoculture of L. vannamei; POLYCUL-SV, polyculture of both species; org.,
organisms.

r 2005 Blackwell Publishing Ltd, Aquaculture Research, 36, 1075^1084

1077

Feeding habits of white and blue shrimp L R Mart| nez-Cordova & E Pen
a-Messina Aquaculture Research, 2005, 36, 1075^1084

Table 2 Mean abundance ( SD) of the main zooplanktonic groups during the experiment in the treatments
Mean  SD
Tr

Weeks

Copepods

MONO-S

1
2
3
4
5
6
7
8
9
10

59
25
407
111
188
384
255
49
98
233












29
135
319
38
89
219
217
16
57
172

25
12
15
24
4
9
7
11
2
4












34
10
17
48
5
11
8
22
3
5

494
351
1355
292
422
441
732
172
284
265












213
128
923
170
323
86
339
105
272
75

275
386
314
53
451
628
191
38
311
357












263
306
451
53
524
606
305
38
390
371

9
3
35
1
0
1
0
0
0
8

1
2
3
4
5
6
7
8
9
10

78
71
31
81
36
46
17
66
24
26












62
40
14
70
13
31
12
58
12
12

1
1
1
7
0
1
1
4
0
1






0
1
1
8

253
260
388
183
141
250
215
119
69
136












129
136
141
30
51
77
99
23
48
36

41
46
70
190
44
63
38
135
31
33












30
50
72
213
32
73
30
174
26
41

5
4
1
0
0
1
0
1
0
0

1
2
3
4
5
6
7
8
9
10

57
68
66
185
44
38
49
128
22
22












35
29
29
150
25
30
33
109
7
9

6
2
1
1
0
2
0
0
0
1

382
342
274
596
184
466
161
356
91
265












89
84
189
394
100
252
123
228
39
162

79
41
37
202
38
196
24
104
25
116












18
26
14
154
21
291
13
65
19
170

16
5
3
0
1
2
2
0
0
1

MONO-V

POLYCUL-SV

Polychaeta

1
0
5
1





11
2
1
2

4

1

Nauplii

Polychaeta larvae

Others

1
 47
1
3

 15
9
3
1

1
1

 23
2
2
1
4
1

1

Values are expressed in thousands of organisms (org.) m .

MONO-S, monoculture of Litopenaeus stylirostris; MONO-V, monoculture of L. vannamei; POLYCUL-SV, polyculture of both species.

Figure 1a^d represent the diet component variations in both shrimp species in the two culture regimes. Not all the components were present in both
species and regimes. Litopenaeus stylirostris in monoculture had a greater diversity in the number of components in its diet, compared with the same species
in polyculture. Litopenaeus vannamei had the same
components in both regimes.
Seven components in the stomach content of the
two species were identied: shrimp exuvia (EXUVIA);
sand grains (ARENA); zooplankton and benthos prey
(PRESAS); unidentied organic matter (MONI); pelleted feed (ARTIF); macroalgae (MACRO) and microalgae (MICRO). The empty portion of the stomachs
studied was called the relative vacuity (Vr). A gradual
decrease in the dierent diet components of these
species was observed from the fth sampling to the
end of the experiment in both regimes.

1078

Table 5 summarizes the proportion of diet components for both species in the two culture regimes. The
most important component for both species, considering the proportion recorded in the stomach contents, was the unidentied organic matter. For L.
vannamei, the proportion of this component was
greater than for L. stylirostris. The component
MACRO was important in few sample weeks. For L.
vannamei, a slightly larger proportion of this component was recorded, reaching 20% in monoculture in
sample week 4. The component ARTIF was only a
small proportion (less than 4%) of the stomach content of L. stylirostris in monoculture, and was not detected for L. vannamei. The component PRESAS had
apparent constant proportions for the two species in
both regimes, although in L. vannamei gut contents,
there were slightly smaller amounts than in L. stylirostris. In monoculture, L. stylirostris showed a notice-

r 2005 Blackwell Publishing Ltd, Aquaculture Research, 36, 1075^1084

Aquaculture Research, 2005, 36, 1075^1084 Feeding habits of white and blue shrimp L R Mart| nez-Cordova & E Pen
a-Messina

Table 3 Mean abundance (  SD) of the main macrozoobenthos groups during the experiment in the treatments
Mean  SD
Tr

Weeks

MONO-S

1
2
3
4
5
6
7
8
9
10

31
493
678
154
62
432
401
216
0
370

1
2
3
4
5
6
7
8
9
10

0
123
463
493
555
0
62
555
216
154

1
2
3
4
5
6
7
8
9
10

0
740
278
432
185
308
185
2251
925
339

MONO-V

POLYCUL-SV

Copepods









Polychaeta

62
439
1276
185
71
433
582
432

 472

Nematods

Insects

0
154
123
370
956
2313
987
216
278
370











185
101
579
722
2775
1204
155
211
142

0
1048
432
771
2529
3515
2652
3824
956
709











2015
370
756
2729
3809
2527
5430
1077
678

0
216
185
1634
771
93
93
278
31
1727











254
236
2239
1132
118
118
408
62
2186

216
339
617
93
0
0
0
216
0
0






101
174
476
395
2443
1075
185
123
62

31
123
463
894
1912
863
863
1233
247
154












62
101
466
476
3105
1406
799
726
142
185

0
216
216
586
4070
2097
1048
401
62
216











274
254
546
5965
2734
1039
582
123
254

93
0
339
216
0
0
93
0
62
0

 118





















247
211
118
476
310
354
142
395
101
790

216
678
308
2251
463
12 920
14 986
1357
709
185












432
498
294
3780
536
16 586
20 733
1126
729
159

0
524
93
493
1295
1819
3084
8326
5581
2159

808
185
439
608
1279
2856
12 484
10 509
2429

31
555
432
278
62
0
0
93
31
0












101
555
363
574






123
409
254
233

0
123
123
278
586
1696
863
93
62
93

1153
274
294
236
467
370
2426
796
599

123
216
216
463
555
524
247
278
123
802











Ostracods











Other

354
420
439
118

 432

 118
 118

 185
 123
62
881
356
395
123

123
62
31
0
0
31
31
123
31
31

 101
 71
 62

185
0
31
0
0
308
185
31
0
62

 123

401
62
62
0
0
31
62
93
31
93

 185
 62







62
62
174
62
62

 62

 370
 370
 62
 71
 484
 71
 71







62
71
185
62
118

Values are expressed in thousands of organisms (org.) m 2.

MONO-S, monoculture of Litopenaeus stylirostris; MONO-V, monoculture of L. vannamei; POLYCUL-SV, polyculture of both species.

Table 4 Global stomach repletion rate (Rsg%), full-stomach index (fr%) and empty stomach index (fv%) in both species and
culture regimes
Monoculture

Polyculture

Litopenaeus stylirostris

L. vannamei

L. stylirostris

Weeks

Rsg%

fr%

fv%

Rsg%

fr%

fv%

Rsg%

fr%

fv%

Rsg%

fr%

fv%

1
2
3
4
5
6
7
8
9
10

41.3
63.1
24.1
57.8
43.7
51.6
71.9
48.4
25.0
7.8

12.5

37.5

12.5

50.0
12.5
12.5
25.0

25.0
37.5

38.1
70.9
86.6
92.2
75.0
73.4
64.7
50.0
47.5
12.5

25.0
50.0
50.0
37.5
25.0
25.0

12.5

12.5
12.5

12.5
12.5

25.0
37.5

42.8
49.1
29.7
55.9
48.4
65.6
21.9
50.6
42.8
41.2

25.0

37.5

12.5

12.5
37.5
50.0

25.0
12.5
25.0

38.1
55.3
35.9
61.9
90.6
47.5
73.4
35.9
40.6
9.4

37.50
12.50
50.00

12.5

25.0

37.5
12.5
50.0

r 2005 Blackwell Publishing Ltd, Aquaculture Research, 36, 1075^1084

L. vannamei

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Feeding habits of white and blue shrimp L R Mart| nez-Cordova & E Pen
a-Messina Aquaculture Research, 2005, 36, 1075^1084

Figure 1 (a) Diet components variation of Litopenaeus stylirostris in monoculture, during the experiment. (b) Diet components variation of L. stylirostris in biculture, during the experiment. (c) Diet components variation of L.s vannamei in
biculture, during the experiment. (d) Diet components variation of L. vannamei in biculture, during the experiment.

Table 5 Components of the stomach content for Litopenaeus vannamei and L. stylirostris in monoculture and polyculture
regimes

Components

MONO-S

VR
EXUVIA
ARENA
PRESAS
MONI
ARTIF
MACRO
MICRO

55.62
0.65
4.79
6.98
27.26
0.92
3.69
0.08










19.48a
1.45a
4.14a
5.90a
10.1a
0.00a
3.25a
0.16a

POLYCUL-SV
(Litopenaeus stylirostris)
54.48
1.45
3.63
6.21
29.78
0
4.45
0







11.74a
2.29a
1.58a
3.50a
8.95ab

 2.99a

MONO-V
36.96
1.88
5.26
4.47
46.04
0
5.34
0







24.41a
3.00a
3.91a
3.20a
17.48b

 6.64a

POLYCUL-SV (L. vannamei)

48.42
2.41
3.12
4.66
38.44
0
2.92
0







22.33a
4.50a
2.15a
4.59a
20.13ab

 2.25a

Mean values  SD (Kruskal^Wallis 95%).

Dierent letters in a row indicate signicant dierences (Po0.05).
MONO-S, monoculture of L. stylirostris; MONO-V, monoculture of L. vannamei; POLYCUL-SV, polyculture of both species.

able increase in prey consumption in the fourth week

of the experiment. The proportion of ARENA was
variable during the experiment and, in some samplings, reached 10% or more of the stomach content
for both species. The presence of the component EXUVIA occurred in the last weeks of the experiment,
starting from the seventh week in polyculture and
the ninth week of monoculture for L. stylirostris. For
L. vannamei, it was found starting from the sixth

1080

week in monoculture and seventh week in polyculture. Microalgae (MICRO) were found in proportions
little more than 1% of the stomach volume in some
samplings for both species.
The production variables for both species in the
two regimes are summarized in Table 6. Growth of L.
vannamei was lower than for L. stylirostris. The rst
grew better in polyculture (10.3  3.4 g) that in
monoculture (9.0  3.8 g). Conversely, L. stylirostris

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Aquaculture Research, 2005, 36, 1075^1084 Feeding habits of white and blue shrimp L R Mart| nez-Cordova & E Pen
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Table 6 Production parameters for Litopenaeus vannamei and L. stylirostris in monoculture and polyculture regimes
Species

Regime

Weight gain (g)

Survival (%)

Production (tonnes ha

L. vannamei

Monoculture
Polyculture
Monoculture
Polyculture

9.03
10.34
16.10
13.43

35.04
41.04
18.08
20.20

0.9485
1.2730
0.8735
0.8140

L. stylirostris

FCR
2.88
2.58
2.99
2.77

FCR, feed conversion ratio.

grew better in monoculture (16.1  4.8 g) than in

polyculture (13.4  4.5 g). Low survival caused a
poor yield for both species, however, L. vannamei
had slightly higher levels. For both species in both regimes, FCR was higher than that reported for commercial purposes (Mart| nez-Cordova 1999). This was
a consequence of a low survival and yields.

Discussion
The large abundance of zooplankton found in this
study, similar or greater than those reported by other
authors (Spanopoulos-Hernandez, Sol-Hernandez,
Zarco-Padron & Franco-Nava 1991; Mart| nez, Barraza
& Pasten1997; Mart| nez-Cordoba et al.1998), suggests
that the fertilization regime was adequate. The greatest density of total macrozoobenthos was found in the
polyculture treatment: 6.9  103  11000 organisms
(org.) m 2. These values are less than those reported
by Barraza-Guardado (1996), who found an abundance of 11  103 org. m 2 in semi-intensive culture
ponds of L. vannamei. The distribution of zoobenthos
is usually in the form of local patches related to the
areas of accumulated organic matter, explaining the
large standard deviations reported here.
Stomach repletion indicies showed higher food
consumption by L. vannamei than by L. stylirostris,
which suggests that the rst species is a more voracious organism or more adaptable to the food present
in the ponds, as suggested by Boddeke (1983). For
both species, a gradual decrease in the stomach repletion rate (Rsg%) was observed, starting from the
fth and seventh week to the end of the experiment.
This can be explained by the decrease of natural food
that is typical for shrimp culture ponds. Further, the
decrease of the water temperature in the ponds over
the course of the study could have had a negative effect on feed intake by the shrimp.
The most important component of the stomach
content for both species, measured by volume, was
organic matter or detritus. For benthic organisms
such as shrimp, detritus constitutes a large propor-

tion of their feed (Muir & Roberts 1982). It has been

noted by many authors that L. vannamei fed on almost anything, including large quantities of organic
matter or detritus in the absence of other food
sources. Litopenaeus stylirostris has been shown to
have restricted nutritional habits (Boddeke 1983);
therefore this species is clearly disadvantaged in
ponds where there may be a limited variety of food
sources compared with L. vannamei. Some reports
support the hypothesis that detritus is the most important food source for penaeid species such as Metapenaeus monoceros, M. japonicus and L. vannamei. The
energy value of detritus has been calculated to be larger that of zooplankton and phytoplankton (Kumari,
Sumitra-Vijayaraghavan, Wafar, Royan & Rajendran
1978). Qasim and Easterson (1974) found that shrimp
have a greater assimilation eciency of estuarine
detritus (93%) than of other food sources. However
some authors report there is no experimental evidence that larger organisms which are metabolically
active and quick growing, like crustaceans, can use
detritus as a primary food source even if this it is microbially enriched (Tenore, Cammen, Finlay & Phillips 1982).
Formulated feed was found in stomach content at
low amounts and only for L. stylirostris in monoculture. Some of what we recorded as ARTIF were identiable pellet fragments. The degree of maceration of
foods by the gastric mill of shrimp can reduce the pellets, microcrustaceans, macroalgae and insects to a
ne mixture. Thus it is probable that a part of the consumed formulated feed was recorded as organic matter. Another possibility is that formulated feed was
consumed by L. vannamei after it has been incorporated to detritus and reached a state of advanced degradation. Reymond and Lagardere (1989) described
the relative role of feed and secondary production in
the diet of M. japonicus. They found that formulated
feed intake did not exceed 4%, an insignicant value
compared with the consumption of other constituents. Cam (1987) and Rollet (1986) found that
approximately 65% of the formulated feed added to
the ponds was not consumed, suggesting that a high

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portion of supplemental feed is not directly consumed by shrimp, but it is indirectly assimilated in diverse links of the trophic chain, particularly through
bacteria (Moriarty, Cook, Bin Hassan & Thanabal
1983; Anderson et al.1987). This assumption has been
demonstrated in more recent studies (Law, Jumars &
Armbrust 2002; Buord, Preston, Nigel, Truong,
Tran, Stuart & Fry 2004). Cam (1987) estimated that
after120 days of culture, only 36% of the organic carbon came from the formulated feed, 43% from secondary productivity and 22% from the unconsumed
feed incorporated into the trophic chain in the ponds.
These results show the necessity of re-evaluating
both the role of supplemental feeding and the role of
natural productivity in shrimp culture.
The EXUVIA in the stomach contents can be from
the internal stomach integument moult. It has been
assumed that penaeid shrimp can digest chitin and
that under extreme conditions can partially consume
their integument. The recording of this component in
the stomach content coincided with a decrease of the
global stomach repletion rate (Rsg%) and with a decline of water temperature in the ponds. This may
generate massive moulting and cause shrimp to stop
feeding and remain buried in the bottom sediment.
Invariably during the experiment, a certain
amount of ARENA was found in the stomach contents of shrimp. Although this component is assumed to possess no nutritional value in penaeid
diets, its presence could suggest a certain importance
in the maceration of food. The presence of sand grains
in the stomach of shrimp is associated with consumption of detritus. In our work, the intake of sand
grains was between 3% and 5% for the two species.
Reymond and Lagardere (1989) found that a small
amount of mineral remains in the stomach content
of M. japonicus.
The presence of prey remains in the gut content of
both species ranged between 4.5% and 7%, with
slightly higher values for L. stylirostris. However, no
signicant dierences were detected between species. Little information exists about prey intake for
these American species. Mart| nez-Cordoba and colleagues (1998) described the presence of copepods,
microcrustacean nauplii and adult copepods in the
stomach contents of L. vannamei. More intensive studies of feeding habits have been carried out for Asian
species. Lanari and colleagues (1989) described 16
components of animal origin in the stomach contents of M. japonicus, with emphasis on the opportunism and predatory activity of this species on
chyronomid insect larvae. This was also reported by

1082

Lanari, Buttazzoni, Tibaldi, Pinosa, Piscitelli and

Carpanese (1985), Gunderman and Poper (1977),
Rubright and colleagues (1981) and Lumare and colleagues (1985) for the same species.
Macroalgae ingestion was important in some sample weeks for both species, but especially for L. vannamei. The ingested species belonged to the genera
Gracilaria and Enteromorpha, which agrees with Mart| nez Cordova (1994) in relation to the stomach
content of L. stylirostris. Some other authors have reported the consumption of macroalgae for farmed
shrimp (Porchas Cornejo, Mart| nez Cordova, Magallon Barajas & Portillo Clark 1999). Few microalgae
were found in the stomachs of the two shrimp species. There is a controversy on the intentional consumption of microalgae by shrimp. It is known that
microalgae play an essential role for nutrition in the
rst larval stages of many shrimp species. However,
in juvenile and adult stages, their importance has
been questioned by some authors, who believe that
the presence of microalgae in the digestive tract is accidental because of the ingestion of detritus. Others
have considered that the contribution of microalgae
becomes important for certain penaeid species (Martinez-Cordova et al. 2003).
The polyculture regime had a dierent eect on the
productive response of the two shrimp species. The
weight gain of L. stylirostris in that regime was significantly lower than in monoculture. The results for L.
vannamei were the opposite.
Our study demonstrates that L. vannamei bases its
feeding almost totally on components associated
with sediment and benthos prey, while L. stylirostris,
with its restricted alimentary characteristic, depended slightly less on detritus and slightly more on
zooplankton prey and pelleted feed. This resulted in
the faster growth of L. stylirostris in monoculture
where the largest zooplankton abundance was recorded. Survival of L. stylirostris was very low during
the experiment. We believe that this was because of
the poor quality of the larvae, which are susceptible
to viral infections. This appeared to be a widespread
problem for larval production of the species in northwest Mexico during the time in which the study was
carried out.
Acknowledgments
We wish to thank CONACYT (The Mexican Council of
Science and Technology) for nancially supporting
this study. Thanks are due to Dr Ellis Glazier for editing the English-language text.

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