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Rearing of misid stages of Penaeus vannamei fed cultured algae of three species
Aq~c~lture, 58 (1936) 139-144 Elsevier Science Publishers B.V., Amsterdam -
139 Printed in The Netherlands
Technical Paper
Rearing of Mysid Stages of Pe~ae~~ va~~a~~~ Fed Cultured Algae of Three Species R~SELBNA SANCHEZ M. ~niuerei~ of Miami, Rosen&e1 S¬ of Marine and Atmos~keric Science, 4600 Rickenb~ker Causeway, Miami, FL 33149 (U.S.A.) Present address: Universidad National Experimental Francisco de Miranda, Centro de Investigaciones Marinas, Plaza Antillana, La Vela de Core, E&do Falcon (Venezuela 1 (Accepted 6 May 1986)
ABSTRACT Srinchez M., R., 1986. Rearing of mysid stages of Penaeus ~a~narne~ fed cultured algae of three species. A~uacu~ture,58: 139-144. Larvae of Penaeus vannamei were reared successfully (53% survival) on an algal diet through the mysid stages without the use of Artemia nauplii. Survival rate and developmental success of larvae fed Isoc~~s~ sp. were superior to those of larvae fed the other algal species tested: Baeteri~trum kya~in~m and Prorocentr~m micans. Differences in percentage survival and developmentaf rate due to the effect of the different diets were significant at P ( 0.01.
INTRODUGTION
Protozoea larvae of penaeids are planktotrophic, As the protozoea metamorphose into myses, their feeding pattern changes from phytoplankton feeding to omnivorous feeding (Heinen, 1976). Mysid stages of penaeids can be reared solely on ph~oplankton but survival is enhanced by animal food ( Hudinaga, 1942). Hudinaga and Kittaka (1967) noted that mysis survival of $8.5% of P. ju~o~ic~ is possible on a diet consisting exclusively of planktonic diatoms. Aquacop (1977) observed that although mysis stages are known to be carnivorous, those of P. monodon could feed on algae up to the postlarval stage. Similarly, Buna Bombeo (1980) reared P. ~ono~on nauplii to postlarvae on ph~oplankton ( ~e~r~e~~~ chuii and C~toceros culcitru~~ . Partly because of rapidly rising costs of A~te~~u eggs in recent years and the accumulating evidence that the nutritional value of Artemiu nauplii and the hatching rate may vary with the locality (Bookhout and Costlow, 1970; Wickins, 1972)) considerable interest has developed in identifying practical, eco-
0 1986 Elsevier Science ~blishe~
B.V.
140
nomical, and nutritional alternatives for Artemia during the mysid stages of penaeid species reared in artificial conditions. There is a need to evaluate the nutritional suitability of readily cultivable algal foods. Three species of alga grown in single-species cultures were used as food for larvae of the white shrimp, Penueus vannamei: Isochrysis sp., Bacteria&rum hyalinum and Prorocentrum micans. These species were selected because of possible differences in their nutritional value related to their classification in the three groups, prymnesiophytes, diatoms and dinoflagellates, respectively. In addition, they differ in size and in mo~holo~, i.e., single cells (Isoch~s~ sp. and P. micans) compared with chain-forming cells (B. hyalinum) . Survival and developmental rates of shrimps were studied during the protozoeal stage, and some batches were reared to the postlarval stage. MATERIALS AND METHODS
Three species of algae: Isochrysis sp. (Tahitian strain, 6pm in diameter), 13. hyalinum (individual cells, 30pm in length, spines not included), and P. micans (32 x 25 pm) were used as food for P. vannamei larvae. The ph~oplankton species were cultured in a modified “f/2” medium (Guillard, 1972). Cultures were harvested during the exponential growth phase and fed to the larvae. Feeding experiments were conducted on a dry weight basis to standardize the addition of different algal species by giving equal amounts of organic matter. This took into account the differences in cell size. Food density was monitored daily by cell counts under the microscope with a hemacytometer. Feeding level was set at 0.0058 mg dry weight/ml. This feeding level was maintained by adding cultured phytoplankton daily. Fluctuation in concentration between additions ranged from 3.80 to 6.50~ lo5 cells/ml for Isochrysis sp., 0.70 to 1.30~ lo* cells/ml for B. hyalinum and 1.06 to 1.70~ lo3 cells/ml for P. micans. About 30% of the total volume of seawater was changed daily, starting from the fourth day of the experiment. Water exchange was made before feeding and after water temperature and salinity had been measured. Nauplius larvae of P. vannamei were obtained from “Intensive Culture Systems”, a privately owned hatchery located on Summerland Key, Florida, and stocked in 5-gallon glass carboys each containing 12 1of filtered seawater provided with aeration, at a density of 50 nauplii per liter. P. vannumei were reared from P 1 to PL 1 ( protozoea 1 to postlarva 1) . Daily estimates of larval population and stages of development were determined by sampling. A total volume of 250 ml was drawn as a sample. Developmental stages were observed under a dissecting microscope and/or compound microscope and compared to the described larval cycles of other penaeids f Wheeler, 1960; Dobkin, 1961; Rodriguez de la Cruz, 1975; Ul-Hassan, 1982 ) .
141
NUIZDEROF DAYS
Fig. 1. Survival rati {mean of two replicates) of Penaeus uu~~~~~ rt3md from nauplius 5 to postiarva 1 on algal diets tQ.00~8 mg dry w~i~~t/mi) .
Rate of development was quantized by means of the growth index for h~vae of Villegas and Kanazawa f 1979 ) s Two replicates were made for each of the treatments arranged in a s~atjal~~ randomized design, The data were subjected to one-way analysis of variance (Sokal and Rohlf? 1981) .
Mean survivsll rates of P. vannumei nauplius 5 to postlarva 1 are shown in Fig. 1. Severe mortality occurred in the populations of larvae fed B. hyalinurn and P. micans during the protozoeal stage, while survival of larvae fed Isochvsis sp. decreased gradually during the protozoeal and mysidstages. Results of the analysis of variance showed that the differences in survival rate due to the effect of different diets were highly significant fP -cO.ODI1. The 95% multiple comparison intervals among pairs of diets by the T-Method ( Sokal and RobIf, L981) showed that all diets were significantly different from each other. Highest percent survival was obtained when larvae were fed Xso&y&s sp. compared with P. ~~cu~$ and B. ~yu~~~u~*
142
7
,*
.I’ i
0
,
=
1110.94
**
.’
i
NUMBER OF DAYS
Fig. 2. Development (mean of two replicates) of Penaeus uannamei reared from nauplius 5 to postlarva 1 on aigal diets (0.0058 mg dry weight/ml).
The effect of each phytoplankton species on the daily mean developmental stage attained by P. vannamei larvae is shown in Fig. 2. On all phytoplankton diets, P. uannamei larvae showed a similar pattern of development for the first day. On the second day, more than 90% of the surviving population fed Isochrysis sp. and P. micuns molted to protozoea 2. In contrast, larvae fed B. hyulinum were considerably delayed in development: it took 5 days for the larval population to completely molt to protozoea 2. These larvae did not survive beyond the fifth day. Clear differences in developmen~l changes of larvae fed ~$oc~~sis sp. and P. micas started on day 3. Larvae fed lsochrysis sp. showed the highest developmental rate and by day 5 more than 50% of the surviving population had developed to mysis 1, while the population fed P. micuns continued at the P 2 stage. Larvae on this diet developed to P 3 but did not survive beyond the seventh day. Results of the analysis of variance showed that the differences in the development rate due to the effect of different diets were significant (P < 0.01) . The 95% multiple comparison intervals among pairs of diets revealed that larvae fed Isochrysis sp. attained the highest developmental rate as compared to B. hyul~n~m and P. micuns.
143 DISCUSSION
Although biochemical analyses for comparisons of the potential nutritive value of the chosen phytoplankton species were not carried out, it seems plausible that differences in survival and developmental rates were due to some extent to the greater ease with which the shrimp ingested the small, naked flagellate, lsochrysis sp. Larvae offered long-chain diatoms and the large dinoflagellate did not survive as long. Spines and other processes may have prevented the larvae from ingesting B. ~y~Zin~~. The armored dinoflagellate, P. ~~c~~s, did not sustain high survival rates of the early protozoeal stages of P. u~~~~e~, while fsochrysis sp. did. The study confirms that larvae of P. uannamei can be reared successfully (53% survival) to PL 1 on an algal diet when fed Isochrysis sp. Of all the chosen phytoplankton species, Isochrysis sp. (UTEX LB 2307) was the easiest to culture, which makes this algal food preferable for the largescale culture of P. uannamei. Semi-continuous cultures of Isochrysis sp., grown in 5-gallon carboys, averaged about 1.18 and 1.22 divisions per day at culture temperatures of 21-25°C. The carboy-cultures yielded a density of about 3.9 x lo6 cells/ml. fsochrysis sp. is nutritionally valuable and may be a useful dietary supplement during the mysid stages of P. ~un~a~ei reared in artificial conditions. ACKNOWLEDGEMENTS
The work reported is based partly on a thesis submitted in partial fulfillment of the requirements for the M.S. degree from the University of Miami. The research was financed by the Universidad National Experimental Francisco de Miranda, Coro, Venezuela. My sincere gratitude goes to Dr. Larry Brand for his guidance at every stage of this study. Special thanks to Drs. Scott Siddall and Nelson Ehrhardt for their assistance in the statistical analyses and to Jesus E. Conde, Centro de Investigaciones Marinas (CIMAR) , Coro, Venezuela, for suggesting improvements to the manuscript.
REFERENCES Aquacop, 1977. Observations sur la maturation et la reproduction en captiviti des crevettes peneida en millieu tropical. In: Third Meeting of the I.C.E.S. Working Group on Mariculture, Brest, France, May 1977. Actes Colloque C.N.E.X.O., 4: 157-178. Bookhout, C.G. and Costlow, J.D., Jr., 1970. Nutritional effects of Artemiafrom different locations on larval development of crabs. Helgol. Wiss. Meeresunters., 20: 435-442. Buna Bombeo, I., 1980. The effects of densities of two food organisms on the rates of survival and
144 development of Penaeus monodon from nauplius VI to postlarvae I. University of the Philippines. 35 pp. ( ms ) . Dobkin, S., 1961. Early development stages of pink shrimp, Penaeus duoraram, from Florida waters. Fish. Bull., 61: 321-349. Guillard, R.R.L., 1975. Culture of phytoplankton for feeding marine invertebrates. In: W. Smith and M. Chanley (Editors), Cultures of Marine Invertebrate Animals. Plenum, New York, pp. 29-60. Heinen, J.M., 1976. An introduction to culture methods for larval and postlarval penaeid shrimps. Proc. World Maricult. Sot., 7: 333-343. Hudinaga, M., 1942. Reproduction, development and rearing of Penaeus japonicus Bate. Jpn. J. Zool., 10(Z): 305-393. Hudinaga, M. and Kit&&a, J., 1967. The large scale pr~uction of the young kuruma prawn, Penaeus ~a~o~~c~ Bate. Information Bulletin on Plank~lo~ in Japan, Commemoration Number of Dr. Y. Matsue’s Sixtieth Birthday, pp. 35-46. Mock, C., Fontaine, C.T. and Revera, D.B., 1980. Improvements in rearing larval penaeid shrimp by the Galveston Laboratory method. In: G. Persoone, P. Sorgeloos, 0. Roels and E. Jaspers (Editors), The Brine Shrimp Artemia, Vol. 3. Universa Press, Wetteren, Belgium, pp. 331-342. Rodriguez de la Cruz, M.C., 1975. Sinopsis biologica de las especies de1 genera Penaeus de1 Pacific0 Mexicano. En: Memorias de1 Simposio sobre Biologfa y Din&mica Poblacional de Camarones, Guaymas, Sonora, August 1976, pp. 282-316. Sokal, R.R. and Rohlf, F.J., 1981. Biometry f 2nd edn). W-H. Freeman, San Francisco, CA, 859 PP. Ul-Hassan, H., 1982. The larval development of Penaew semisukatus De Haan 1850 (Decapoda, Penaeidae) reared in the laboratory. J. Plankton Res., 4( 1) : 3-17. Villegas, C.T. and Kanazawa, A., 1979. Relationship between diet composition and growth rate of the zoeal and mysis stages of Penaeus japonicus Bate. Southeast Asian Fisheries Development Center, Q. Res. Rep., II( 2): 24-29. Wheeler, R.S., 1960. Experiments rearing shrimp larvae. U.S. Fish. Wildl. Ser. Circ., 92: 7-8. Wickins, J.F., 1972. The food value of the brine shrimp Artemia salina L., to larvae of the prawn, Pataemon serrates Pennant. J. Exp. Mar. Biol. Ecol., 10: 151-170.
139 Printed in The Netherlands
Technical Paper
Rearing of Mysid Stages of Pe~ae~~ va~~a~~~ Fed Cultured Algae of Three Species R~SELBNA SANCHEZ M. ~niuerei~ of Miami, Rosen&e1 S¬ of Marine and Atmos~keric Science, 4600 Rickenb~ker Causeway, Miami, FL 33149 (U.S.A.) Present address: Universidad National Experimental Francisco de Miranda, Centro de Investigaciones Marinas, Plaza Antillana, La Vela de Core, E&do Falcon (Venezuela 1 (Accepted 6 May 1986)
ABSTRACT Srinchez M., R., 1986. Rearing of mysid stages of Penaeus ~a~narne~ fed cultured algae of three species. A~uacu~ture,58: 139-144. Larvae of Penaeus vannamei were reared successfully (53% survival) on an algal diet through the mysid stages without the use of Artemia nauplii. Survival rate and developmental success of larvae fed Isoc~~s~ sp. were superior to those of larvae fed the other algal species tested: Baeteri~trum kya~in~m and Prorocentr~m micans. Differences in percentage survival and developmentaf rate due to the effect of the different diets were significant at P ( 0.01.
INTRODUGTION
Protozoea larvae of penaeids are planktotrophic, As the protozoea metamorphose into myses, their feeding pattern changes from phytoplankton feeding to omnivorous feeding (Heinen, 1976). Mysid stages of penaeids can be reared solely on ph~oplankton but survival is enhanced by animal food ( Hudinaga, 1942). Hudinaga and Kittaka (1967) noted that mysis survival of $8.5% of P. ju~o~ic~ is possible on a diet consisting exclusively of planktonic diatoms. Aquacop (1977) observed that although mysis stages are known to be carnivorous, those of P. monodon could feed on algae up to the postlarval stage. Similarly, Buna Bombeo (1980) reared P. ~ono~on nauplii to postlarvae on ph~oplankton ( ~e~r~e~~~ chuii and C~toceros culcitru~~ . Partly because of rapidly rising costs of A~te~~u eggs in recent years and the accumulating evidence that the nutritional value of Artemiu nauplii and the hatching rate may vary with the locality (Bookhout and Costlow, 1970; Wickins, 1972)) considerable interest has developed in identifying practical, eco-
0 1986 Elsevier Science ~blishe~
B.V.
140
nomical, and nutritional alternatives for Artemia during the mysid stages of penaeid species reared in artificial conditions. There is a need to evaluate the nutritional suitability of readily cultivable algal foods. Three species of alga grown in single-species cultures were used as food for larvae of the white shrimp, Penueus vannamei: Isochrysis sp., Bacteria&rum hyalinum and Prorocentrum micans. These species were selected because of possible differences in their nutritional value related to their classification in the three groups, prymnesiophytes, diatoms and dinoflagellates, respectively. In addition, they differ in size and in mo~holo~, i.e., single cells (Isoch~s~ sp. and P. micans) compared with chain-forming cells (B. hyalinum) . Survival and developmental rates of shrimps were studied during the protozoeal stage, and some batches were reared to the postlarval stage. MATERIALS AND METHODS
Three species of algae: Isochrysis sp. (Tahitian strain, 6pm in diameter), 13. hyalinum (individual cells, 30pm in length, spines not included), and P. micans (32 x 25 pm) were used as food for P. vannamei larvae. The ph~oplankton species were cultured in a modified “f/2” medium (Guillard, 1972). Cultures were harvested during the exponential growth phase and fed to the larvae. Feeding experiments were conducted on a dry weight basis to standardize the addition of different algal species by giving equal amounts of organic matter. This took into account the differences in cell size. Food density was monitored daily by cell counts under the microscope with a hemacytometer. Feeding level was set at 0.0058 mg dry weight/ml. This feeding level was maintained by adding cultured phytoplankton daily. Fluctuation in concentration between additions ranged from 3.80 to 6.50~ lo5 cells/ml for Isochrysis sp., 0.70 to 1.30~ lo* cells/ml for B. hyalinum and 1.06 to 1.70~ lo3 cells/ml for P. micans. About 30% of the total volume of seawater was changed daily, starting from the fourth day of the experiment. Water exchange was made before feeding and after water temperature and salinity had been measured. Nauplius larvae of P. vannamei were obtained from “Intensive Culture Systems”, a privately owned hatchery located on Summerland Key, Florida, and stocked in 5-gallon glass carboys each containing 12 1of filtered seawater provided with aeration, at a density of 50 nauplii per liter. P. vannumei were reared from P 1 to PL 1 ( protozoea 1 to postlarva 1) . Daily estimates of larval population and stages of development were determined by sampling. A total volume of 250 ml was drawn as a sample. Developmental stages were observed under a dissecting microscope and/or compound microscope and compared to the described larval cycles of other penaeids f Wheeler, 1960; Dobkin, 1961; Rodriguez de la Cruz, 1975; Ul-Hassan, 1982 ) .
141
NUIZDEROF DAYS
Fig. 1. Survival rati {mean of two replicates) of Penaeus uu~~~~~ rt3md from nauplius 5 to postiarva 1 on algal diets tQ.00~8 mg dry w~i~~t/mi) .
Rate of development was quantized by means of the growth index for h~vae of Villegas and Kanazawa f 1979 ) s Two replicates were made for each of the treatments arranged in a s~atjal~~ randomized design, The data were subjected to one-way analysis of variance (Sokal and Rohlf? 1981) .
Mean survivsll rates of P. vannumei nauplius 5 to postlarva 1 are shown in Fig. 1. Severe mortality occurred in the populations of larvae fed B. hyalinurn and P. micans during the protozoeal stage, while survival of larvae fed Isochvsis sp. decreased gradually during the protozoeal and mysidstages. Results of the analysis of variance showed that the differences in survival rate due to the effect of different diets were highly significant fP -cO.ODI1. The 95% multiple comparison intervals among pairs of diets by the T-Method ( Sokal and RobIf, L981) showed that all diets were significantly different from each other. Highest percent survival was obtained when larvae were fed Xso&y&s sp. compared with P. ~~cu~$ and B. ~yu~~~u~*
142
7
,*
.I’ i
0
,
=
1110.94
**
.’
i
NUMBER OF DAYS
Fig. 2. Development (mean of two replicates) of Penaeus uannamei reared from nauplius 5 to postlarva 1 on aigal diets (0.0058 mg dry weight/ml).
The effect of each phytoplankton species on the daily mean developmental stage attained by P. vannamei larvae is shown in Fig. 2. On all phytoplankton diets, P. uannamei larvae showed a similar pattern of development for the first day. On the second day, more than 90% of the surviving population fed Isochrysis sp. and P. micuns molted to protozoea 2. In contrast, larvae fed B. hyulinum were considerably delayed in development: it took 5 days for the larval population to completely molt to protozoea 2. These larvae did not survive beyond the fifth day. Clear differences in developmen~l changes of larvae fed ~$oc~~sis sp. and P. micas started on day 3. Larvae fed lsochrysis sp. showed the highest developmental rate and by day 5 more than 50% of the surviving population had developed to mysis 1, while the population fed P. micuns continued at the P 2 stage. Larvae on this diet developed to P 3 but did not survive beyond the seventh day. Results of the analysis of variance showed that the differences in the development rate due to the effect of different diets were significant (P < 0.01) . The 95% multiple comparison intervals among pairs of diets revealed that larvae fed Isochrysis sp. attained the highest developmental rate as compared to B. hyul~n~m and P. micuns.
143 DISCUSSION
Although biochemical analyses for comparisons of the potential nutritive value of the chosen phytoplankton species were not carried out, it seems plausible that differences in survival and developmental rates were due to some extent to the greater ease with which the shrimp ingested the small, naked flagellate, lsochrysis sp. Larvae offered long-chain diatoms and the large dinoflagellate did not survive as long. Spines and other processes may have prevented the larvae from ingesting B. ~y~Zin~~. The armored dinoflagellate, P. ~~c~~s, did not sustain high survival rates of the early protozoeal stages of P. u~~~~e~, while fsochrysis sp. did. The study confirms that larvae of P. uannamei can be reared successfully (53% survival) to PL 1 on an algal diet when fed Isochrysis sp. Of all the chosen phytoplankton species, Isochrysis sp. (UTEX LB 2307) was the easiest to culture, which makes this algal food preferable for the largescale culture of P. uannamei. Semi-continuous cultures of Isochrysis sp., grown in 5-gallon carboys, averaged about 1.18 and 1.22 divisions per day at culture temperatures of 21-25°C. The carboy-cultures yielded a density of about 3.9 x lo6 cells/ml. fsochrysis sp. is nutritionally valuable and may be a useful dietary supplement during the mysid stages of P. ~un~a~ei reared in artificial conditions. ACKNOWLEDGEMENTS
The work reported is based partly on a thesis submitted in partial fulfillment of the requirements for the M.S. degree from the University of Miami. The research was financed by the Universidad National Experimental Francisco de Miranda, Coro, Venezuela. My sincere gratitude goes to Dr. Larry Brand for his guidance at every stage of this study. Special thanks to Drs. Scott Siddall and Nelson Ehrhardt for their assistance in the statistical analyses and to Jesus E. Conde, Centro de Investigaciones Marinas (CIMAR) , Coro, Venezuela, for suggesting improvements to the manuscript.
REFERENCES Aquacop, 1977. Observations sur la maturation et la reproduction en captiviti des crevettes peneida en millieu tropical. In: Third Meeting of the I.C.E.S. Working Group on Mariculture, Brest, France, May 1977. Actes Colloque C.N.E.X.O., 4: 157-178. Bookhout, C.G. and Costlow, J.D., Jr., 1970. Nutritional effects of Artemiafrom different locations on larval development of crabs. Helgol. Wiss. Meeresunters., 20: 435-442. Buna Bombeo, I., 1980. The effects of densities of two food organisms on the rates of survival and
144 development of Penaeus monodon from nauplius VI to postlarvae I. University of the Philippines. 35 pp. ( ms ) . Dobkin, S., 1961. Early development stages of pink shrimp, Penaeus duoraram, from Florida waters. Fish. Bull., 61: 321-349. Guillard, R.R.L., 1975. Culture of phytoplankton for feeding marine invertebrates. In: W. Smith and M. Chanley (Editors), Cultures of Marine Invertebrate Animals. Plenum, New York, pp. 29-60. Heinen, J.M., 1976. An introduction to culture methods for larval and postlarval penaeid shrimps. Proc. World Maricult. Sot., 7: 333-343. Hudinaga, M., 1942. Reproduction, development and rearing of Penaeus japonicus Bate. Jpn. J. Zool., 10(Z): 305-393. Hudinaga, M. and Kit&&a, J., 1967. The large scale pr~uction of the young kuruma prawn, Penaeus ~a~o~~c~ Bate. Information Bulletin on Plank~lo~ in Japan, Commemoration Number of Dr. Y. Matsue’s Sixtieth Birthday, pp. 35-46. Mock, C., Fontaine, C.T. and Revera, D.B., 1980. Improvements in rearing larval penaeid shrimp by the Galveston Laboratory method. In: G. Persoone, P. Sorgeloos, 0. Roels and E. Jaspers (Editors), The Brine Shrimp Artemia, Vol. 3. Universa Press, Wetteren, Belgium, pp. 331-342. Rodriguez de la Cruz, M.C., 1975. Sinopsis biologica de las especies de1 genera Penaeus de1 Pacific0 Mexicano. En: Memorias de1 Simposio sobre Biologfa y Din&mica Poblacional de Camarones, Guaymas, Sonora, August 1976, pp. 282-316. Sokal, R.R. and Rohlf, F.J., 1981. Biometry f 2nd edn). W-H. Freeman, San Francisco, CA, 859 PP. Ul-Hassan, H., 1982. The larval development of Penaew semisukatus De Haan 1850 (Decapoda, Penaeidae) reared in the laboratory. J. Plankton Res., 4( 1) : 3-17. Villegas, C.T. and Kanazawa, A., 1979. Relationship between diet composition and growth rate of the zoeal and mysis stages of Penaeus japonicus Bate. Southeast Asian Fisheries Development Center, Q. Res. Rep., II( 2): 24-29. Wheeler, R.S., 1960. Experiments rearing shrimp larvae. U.S. Fish. Wildl. Ser. Circ., 92: 7-8. Wickins, J.F., 1972. The food value of the brine shrimp Artemia salina L., to larvae of the prawn, Pataemon serrates Pennant. J. Exp. Mar. Biol. Ecol., 10: 151-170.