Laboratory Culture of the Caspian Sea Calanoid Copepod Acartia clausi (Giesbrecht, 1889) at Different Salinity Levels

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1 World Journal of Fish and Marine Sciences 3 (6): , 20 ISSN IDOSI Publications, 20 Laboratory Culture of e Caspian Sea Calanoid Copepod Acartia clausi (Giesbrecht, 889) at Different Salinity Levels 2 Esmaeili Fereidouni Abolghasem, Shaygan Majid, 3 2 Agh Naser, Ouraji Hossein and Masoudi Asil Shima Fisheries Department, Faculty of Animal Sciences and Fisheries, Sari Agricultural Sciences and Natural Resources University (SANRU), Sari, Iran 2 Faculty of Animal Sciences and Fisheries, Sari Agricultural Sciences and Natural Resources University, Sari, Iran 3 Artemia and Aquatic Animals Research Institute, Urmia University, Urmia, 5753, Iran Abstract: The Caspian Sea calanoid copepod, Acartia clausi, has good potential for mass culture as a live feed for mariculture. This study was carried out to investigate e effects of salinity on density of A. clausi at four salinities of 3, 20, 35 and 45 in 5 L carboys container and mass culture at two salinities of 3 and 35 in 00 L fiberglass culture tanks over a two weeks period. A. clausi was cultured at temperature C and fed wi two algal diets including Isochrysis galbana and Chaetoceros calcitrans at food concentration cells mlg. Results showed at salinity had significant effects on density in 5 L container. Wi an initial stocking density of 5 adults LG, maximum and minimum mean density reached to 435.3±75.05 and 03.96±2.56 individuals LG at salinities of 3 and 45 wi significant (P$0.05) differences between treatments, respectively. Population density was not significant at 3 and 35 in 5 L containers (P#0.05). In 00 L tanks wi an initial stocking density of 8-20 adults LG, e mean density was not significant (P#0.05) at two salinities of 3 and 35. Total mean copepod production reached to densities of 048.5±86.4 and ±270.7 individuals LG wi e dominance of nauplius stages at salinities of 3 and 35, respectively. Maximum density was recorded 3437 individuals LG (3267 nauplii, 73 copepodites (C-5) and 97 adults LG ) at salinity of 35. The results of is study recommended at A. clausi is a euryhaline species and can tolerate different salinities in culture conditions; however, it is better to be cultured at salinity of 35 for maximum population density grow. Key words: Copepod % Salinity % Acartia clausi % Density % Caspian Sea % Live Feed INTRODUCTION vitamins (C and E), small size, digestibility and eir swimming motion [2-4], which are essential for larval Copepods constitute a major part of e diet of fish survival, grow, digestion and metamorphosis [5-8]. larvae in e natural pelagic food chain; in which, e Different studies showed at copepods diets show more ree main orders including Calanoida, Harpacticoida significant benefits to increase larval marine fish grow and Cyclopoida have each been investigated for eir and development better an rotifers Brachionus spp. and suitability for larval and juvenile fish at each order has Artemia [6, 9-6]. Despite ese findings, enriched rotifers its advantages and disadvantages []. and Artemia will probably continue to be e live feeds of In aquaculture, copepods are generally considered to choice in commercial hatcheries [5-7]. be nutritionally superior live feeds, as ey are a valuable In general, copepods are difficult to culture at source of protein, lipid (especially highly unsaturated sufficient densities to be economically efficient on a fatty acids contents, 20:5n-3 and 22:6n-3), carbohydrates, commercial scale, because ey require high water enzymes (amylase, protease, exonuclease and esterase), volumes for cultivation in captivity and is is perceived Corresponding Auor: Esmaeili Fereidouni, Fisheries Department, Faculty of Animal Sciences and Fisheries, Sari Agricultural Sciences and Natural Resources University (SANRU), P.O.Box. 578, Sari, Iran. 590

2 World J. Fish & Marine Sci., 3 (6): , 20 to be too expensive and unreliable for most intensive samples were immediately transported to e laboratory hatcheries. Therefore, only a few species of copepods and oroughly rinsed to reduce contamination by have been successfully reared at near commercial scale in unwanted organisms. After rinsing, e zooplanktons extensive systems. Most copepod rearing trials have been were screened to isolate e size fraction containing small scale and lasted only a few weeks or mons [8,8]. predominantly adult copepods of A. clausi [2]. Adults According to e review by [6,8], a reliable system for e (75-00 individuals LG ) were divided randomly into four continuous large-scale indoors intensive culture of parts (each salinity level), which were reared wi A. calanoid, harpacticoid or cyclopoid copepods has not yet clausi in five L beakers. Before e experiments, e five been developed. groups of adult copepods were acclimated gradually from The most frequently cultured calanoid species belong 3 to 20, 35 and 45 and increased five units per day to e genera found in coastal waters, such as ose of during 2 to 6 days period [23]. Healy adults (survival e genera Acartia, Centropages, Eurytemora and Temora. rates approximately were % at different salinities) These copepods are small, wi relatively short generation from e groups were selected for e experiments. times, a wide ermal and salinity tolerance and are easily Various salinity conditions were obtained by diluting adaptable to laboratory conditions and when reared in meod wi e salinity of 55 obtained from e outdoor multispecies cultures, ey tend to dominate wi Gomishan Shrimp Center ponds (Golestan province) wi time [9]. Calanoid copepods of e genus Acartia is dechlorinated freshwater. Salinity was measured by a subtropical and temperate coastal marine, estuarine areas hand refractometer. and semi-enclosed systems wi a cosmopolitan Then, e copepods were separately cultured under distribution [20-22]. Acartia clausi is a free-spawning wi four salinity levels (3, 20, 35 and 45 ) in 5 L carboys more abundant outside e estuary and shows maximum containers and mass cultured under two salinity levels (3 densities during spring-autumn in e Caspian Sea. This and 35 ) in 00 L fiberglass tanks over two weeks species was acclimated to salinity of 3 in e souern culture period. The copepods fed daily wi a mixture of waters of e Caspian Sea. two algal diets including I. galbana and C. calcitrans in a Salinity is one of e most important environmental 3 ratio of : to give a final density at concentration 40 0 parameters affecting e seasonal and spatial distribution cells mlg at measured by haemocytometer. Copepods of marine copepods in e wild and can affect e were cultured at C and 2h L: 2h D (Light: Dark) spawning, incubation, survival rate, grow, respiration photoperiod. Aeration was provided for small and large and subrogation of e dominant species in nature [23]. experimental treatments. For 5 L containers, e aeration Few studies, however, exist on e density of A. clausi was provided centrally near e bottom of e carboys, and its mass culture for aquaculture purposes in using a glass pipette wi an aperture of.5 mm controlled laboratory or hatchery conditions at different salinities. to give a bubble of air one to ree times per second. For Therefore, is paper discusses wi e aim to study e 00 L tanks, aeration was provided by an air stone and its effects of different salinity levels on density of A. clausi intensity was strong enough to circulate e water in e in laboratory conditions. tanks wiout causing excessive turbulence. Oxygen concentrations were regulated at mg LG. For water MATERIALS AND METHODS exchange, % of water volume was approximately exchanged every ree days using a siphon wi a 20 µm Algal Culture: Microalgal cultures used for experiments mesh attached to e end and new seawater at had been were two algal diets including Isochrysis galbana pre-adjusted to desired salinity was added [2]. For all (Prymnesiophyceae) and Chaetoceros calcitrans experiments, ree replicates were set up for each (Bacillariophyceae) at batch cultured in e SANRU treatment. ï phycolab laboratory at 26 C, 4L: 0D (Light: Dark) The copepod densities were sampled every day in photoperiod, 5,000-6,000 Lux illumination and fertilized 5 L and every oer day in 00 L cultures, respectively. wi f 2 medium at different salinities i.e., 3, 20, 35 and 45 After collection, clove oil was used to immobilize e [24]. copepods before ey were counted using a stereo microscope (Model, Nikon-SMZ 500-Japan). Total Copepod Culture: Adult A. clausi were collected using counts of representative samples from e condensed 200-µm mesh size plankton nets from norern coast populations were repeated ree times for each treatment (Mahmoud Abad) of e Caspian Sea (3 ). The and size group, recording e proportion of e population 59

3 World J. Fish & Marine Sci., 3 (6): , 20 in each life stages (egg and nauplius, copepodite and RESULTS adult). Therefore, e population densities of copepods were estimated from counting e total number of The density of adult, copepodites, egg and nauplii copepods in multiple ree subsamples and e numbers and total copepods of all stages were compared between of nauplius instars (NI-N VI ), copepodid instars (CI-C V) and treatments. In 5 L containers, e maximum density of adults (C VI) recorded [5]. copepod was attained during 0 to 3 days of culture at different salinities. It was noticed at e copepod Statistical Analysis: Prior to analysis, e data were density decreased from e 4 days onwards (Fig., 2 arcsin transformed for homogeneity and analyzed by one and 3). There were no significant differences in total way ANOVA. When a significant difference (P#0.05) was copepod density between salinities of 3 and 35 found, Duncan's multiple comparisons test was used to (P$0.05); however, e observed differences between discern significant differences between treatments. All ese treatments were significant (P#0.05) compared to statistical analyses were conducted using SPSS program e 20 and 45 (Table ). Maximum and minimum of total (Version 7). Data are presented as mean ± standard error copepod density per liter were recorded at 3 and 45 (SE). (P#0.05), respectively. Total copepod density was higher Y = 52.6 X R2 = ppt Y = X R 2 = ppt Mean density / lit Mean density / lit Culture days Culture days Fig. : Total production (Mean density per L ± SE) of copepod, A. clausi, at different salinity levels over two weeks culture in 5 L container (Dashed line represented regression linear). 592

4 Mean density / lit World J. Fish & Marine Sci., 3 (6): , at 2 week compared to e week in all treatments (Table ), however, an opposite trend has been observed in nauplii percentage in all 800 treatments. Moreover, during 5 days operation, e 700 systems produced an average of 663 to 6962 individuals 600 LG of copepod at salinity of 45 and 3, respectively (P#0.05). The maximum density was recorded individuals LG at 3 (Table ); however, higher regression coefficient (R ) observed at 3 and (P$0.05) (R =75.69 for 3 and R =82.80 for 35 ) 300 (Fig. ). 200 In 00 L tanks, ere were no significant differences 00 in total copepod density between two salinities levels (P$0.05). Mean copepod density (wi initial stocking 0 density of 8-20 adults LG ) reached to 048 and individuals LG at salinities of 3 and 35, respectively. Culture days Maximum density were recorded 3437 and 2248 Fig. 2: The comparisons of total production (Mean individuals LG at 35 and 3, respectively (Table 2). density per L) of copepod, A. clausi, at different After two weeks of experiment, e produced biomass of salinity levels over two weeks culture in 5 L nauplius was relatively higher (P#0.05) in second week in container. comparison wi e first week at different salinities ppt Nauplii C-5 Adult (C6) Mean density / lit Culture days Fig. 3: Population composition of different life stages (nauplii, copepodites and adults) of copepod, A. clausi, from an initial stocking density of 5 adults LG at different salinities over two weeks culture in 5 L container. 593

5 World J. Fish & Marine Sci., 3 (6): , 20 Table : Total production (Mean density per L ± SE) of copepod, A. clausi at different salinities in 5 L containers over two weeks culture period Salinity ( ) Parameters a b a c Total organism per L 435.3± ± ± ±2.56 a a a b Mean density per L in first week 29.29± ± ± ±9.92 a b a c Mean density per L in second week ± ± ± ±5.20 a a a b Nauplius percentage in first week 83.32± ± ± ±7.37 a b b b Nauplius percentage in second week 76.90± ± ± ±4.62 Min-Max density per L Table 2: Total production (Mean density per L ± SE) of copepod, A. clausi, at two salinities of 3 and 35 in 00 L tanks over a two weeks period Salinity ( ) Parameters 3 35 a Total organism per L 048.5± ± a Mean density per L in first week 670.4±25.29 a 839±8.39 a Mean density per L in second week 520.3± a ± a a a Nauplius percentage in first week 85.2± ±4.07 a a Nauplius percentage in second week 9.37± ±2.54 Min-Max density per L Table 3: Culture meods and productivity for calanoid copepods used as a live feed for marine fish larviculture [,6,8] Species Culture size (L) Densities Productivity Culture period Food conditions Ref. Acartia tonsa /L 2-75 nauplii adultg 6 days Natural phytoplankton, Extensive [4] Acartia tonsa /L eggs LG 28 days Rhodomonas baltica Isochrysis galbana [36] Gladioferens imparipes 500, Automated, batch culture Stocked at 000 nauplii LG 878 nauplii.lg.dg 420 days Rhodomonas baltica Isochrysis galbana [40] 3 Acartia clausi 00, Recirculation - #40 per L 4 mons Rhodomonas baltica (50 0 cells mlg ) Isochrysis galbana 3 (50 0 cells mlg ) [34] 3 Acartia tonsa 00, Recirculation - #40 per L 0 mons Rhodomonas baltica (50 0 cells mlg ) 3 Isochrysis galbana (50 0 cells mlg ) [34] Acartia tonsa 890, Circular outdoor tanks per L 6 mons Natural phytoplankton blooms [4] Acartia clausi adults LG year Tetraselmis suecica [35] Acartia clausi + Tisbe furcata 40, Multiple generations individuals LG Tetraselmis suecica [35] mixed species - Eurytemora affinis individuals LG - Nanochloris sp. [47] Temora longicornis adults LG 2 mons Tetraselmis suecica [35] 4 Temora longicornis 00, Multiple generations - #40 per L - Rhodomonas baltica (2-4 0 cells mlg ) Parvacalanus 4 Isochrysis galbana (3-8 0 cells mlg ) [48] crassirostris individuals LG - Isochrysis/Rhodomonas/Tetraselmis/Heterocapsa [42] Acartia clausi 5,Batch culture Initial stocked at organism LG, Isochrysis galbana Chaetoceros calcitrans The 5 female LG Max:,008 individuals 4 (: ratio; 4 0 cells mlg ) present LG at 3 2 weeks study Acartia clausi 00,Batch culture Initial stocked at Isochrysis galbana Chaetoceros calcitrans The 8-20 female LG organism LG, 4 (: ratio; 4 0 cells mlg ) present Max: 3,437 individuals LG at 35 2 weeks study 594

6 World J. Fish & Marine Sci., 3 (6): , Nauplii C-5 Adult Sum Linear (Sum) 3 ppt Y = X R 2 = Mean density / lit Culture days Fig. 4: Total production (Mean density per L ± SE) and population composition of different life stages (nauplii, copepodites and adults) of A. clausi at two salinities of 3 and 35 over a two weeks culture in 00 L tanks (Dashed line represented regression linear). The nauplius densities of higher an 000 individuals LG laboratory or field conditions [32, 29]. These data are were achieved on day of fif of culture at two above higher an e egg production rate of e Caspian Sea mentioned salinity levels; however, higher densities were copepod, A. clausi. [33] found at wiin e ree observed at 35 during 6 to days of culture; salinities levels (3, 20 and 30 ), maximum egg whereas, in terms of 3, similar densities achieved production of A. clausi was obtained at 30 ; however, during 8 to days of experiments (Fig. 4). salinity had no significant effect on egg production rate, which remained at 6-9 egg.femaleg.dayg at salinity of 3 DISCUSSION and 30, respectively. Alough A. clausi completely adapted to e Calanoid copepods are often well adapted to Caspian Sea brackish water (3 ), our results clearly cope wi seasonal fluctuations in salinity variations revealed at it is a euryhaline species and can tolerate under natural conditions. Maximizing copepod salinity fluctuations and even reproduce in such a productivity is e major goal for aquaculture condition. An increase in salinity caused a sharp purposes and salinity conditions used for culture reduction in nauplius percentage and increased e nd are likely to affect productivity [20]. It is often difficult copepodite (C I-V) and adult density after 2 weeks, kept in to define salinity reshold for estuarine species, as 5 L carboys (apart from 45 treatment) (Fig. 2). As it different species may adapt to different salinities at was expected, higher densities were recorded in 00 L various stages of eir life history due to eir habitat tanks at two salinity levels of 3 and 35 (Fig. 4). variations and it is likely to be species specific, or life- Salinity had no significant effects in itself on stage specific [20,25]. copepod production at two salinities 3 and 35 in Fecundity of copepods is influenced by e density 00 L tanks; however, e salinity of 35 had better of breeding females and nauplii, food quality and results in final copepod production over two weeks quantity, temperature, salinity, turbulence, stocking culture. According to e results of e present study, density, cannibalism of nauplii by adult and higher assuming no culture harvesting, a batch culture of 5 copepodite stages, culture tank size and shape, water days period has been found to be desirable under quality, sex ratio and female longevity [26-3]. Free- laboratory conditions and sufficient for obtaining spawning calanoid copepods such as various Acartia maximum copepods densities for A. clausi. It seems species may produce between -50 eggs.femaleg.dayg, at A. clausi had a development period of 8-0 days at producing a total of up 200 from one single spawning in C in our experiments. 595

7 World J. Fish & Marine Sci., 3 (6): , 20 The obtained densities in our study were higher an [34] and [35]. [34] obtained a density lower an 40 individuals LG at a 4 mons period in 00 L recirculation systems for A. clausi at fed twice weekly wi Rhodomonas baltica and Isochrysis galbana at algal 5 density 0 cells mlg in a ratio :. Also, [35] cultured e same species in one year period in 20 L carboys systems and obtained densities approximately adults LG wi Tetraselmis suecica as a sole diet (mean density in 5 L carboy culture was 405 individuals LG at 35 in e present study). Several attempts to mass-culture copepods in intensive systems have been undertaken wi varying success and have resulted in e development of different systems for particular species of copepods. Alough Acartia species have been cultured successfully for many generations in e laboratory [36] and in large outdoor tanks [37], e low densities likely make calanoids inappropriate for intensive or super-intensive mass cultivation (Table 3). The mass culture of Acartia spp. is related to e available water volume, raer an e available substrate area as in harpacticoids [27]. The different obtained densities in our study can be possibly attributed to different water volume for culture of A. clausi in 5 and 00 L cultures, which is in agreement wi e results of some oer researchers [,36]. The maximum density of A. clausi was 3437 individuals LG (3267 nauplii, 73 copepodids (C I-V) and 97 adult per L) at 35 in 00 L tanks in our experiment, which was lower an e production of calanoid copepod, Acartia souwelli and A. centrura wi e maximum daily production of 485 nauplii, 245 copepodids and,285 adults LG for A. souwelli and 3547 nauplii, 74 copepodids and,42 adults LG in A. centrura during a 5 days period in 25 L tanks [38]. These discrepancies may be attributed to e differences in species, culture meods, culture size, initial stocking density, productivity and food quality and quantity. [38] fed e copepods at a ratio of cells mlg which were higher an our experiments (40000 cells mlg ). Our production figures exceed ose reported for temperate Acartia species. For example, e maximum densities reported for Acartia tsuensis cultured in outdoor tanks were 36 nauplii, 588 copepodids and 280 adults LG [39]. Mean densities of lower an 2000 individuals LG also obtained in an 8-days cycle for Acartia spp. in Darwin center marine fish [5]. However, a daily productivity of 95,000 eggs correspond to 500 nauplii LG wi 45 percent hatching success reported for A. tonsa in 200 L culture tanks [36]. Also, over 7 weeks of operation on Acartia spp. (Australian strain), [2] started wi an inoculums of around adults and copepodites LG, e culture an average produced around 2,000 nauplii, 750 copepodites and 300 adults LG after 7 days (maximum output was,00 copepodids and 729 adults LG ) and a mean peak density of nauplii was almost 2,000 individuals LG (e highest was 5,50 individuals LG ) in,000 L tanks; which were higher an our data about A. clausi. They noted at an 8-day culture cycle seems an appropriate period for mass culture of Acartia ï spp. wi a generation time of 5-7 days at C, and 20 0 cells LG of ree algal diets including Isochrysis, Rhodomonas and Tetraselmis [2]. [40] were able to produce 878 nauplii LG dayg of e calanoid species, Gladioferens imparipes in 500 L automated batch cultures for 420 days. Most calanoid copepods can be grown at densities of only adults lg [4, 36]. [27] discussed possible causes of sex ratio regulation (e female percentage). Crowding may cause changes in e sex ratio, so at low female percentages occurring at high stocking densities. Also, cannibalism of nauplii by adults and later stage copepodids is a problem when trying to maintain high densities of nauplii in copepods cultures [33,36-37]. However, obtained densities in Parvocalanus crassirostris stock cultures have been averaged about 7 nauplii and 3 adult/late stages copepodites per ml after nine days in 20 L culture tanks [5]. Cannibalism may also be a potential contributing factor to e sharp decline in nauplii numbers from e days onwards in 00 L tanks in e current study. Wiin e population in 5 L carboys, ere were some differences in e distribution of e various lifestages (nauplii, copepodites and adults) of A. clausi cultured at different salinities. At e lower salinities (3 and 20 ), a relatively higher proportion of e population was at e nauplius stage, while ere was a more even distribution of different life-stages at e higher salinity of 35 (Fig. 3). These could be due to e necessity of calanoid copepods to higher volume water for maximum egg and nauplius production. It has already been stated at fecundity decreases wi increasing density; in which, population density can affect population grow by modulating survival, development and fecundity [42]. It has been suggested at complex chemical compounds may be produced by e animals as a result of crowding, allowing em to perceive and respond to different crowding levels [27]. Alternatively, it has also been hypoesized at direct close encounter may change e behavior of copepods and even eir development. 596

8 World J. Fish & Marine Sci., 3 (6): , 20 The optimal food concentrations for culture of 2. Shields, R.J., J.G. Bell, F.S. Luizi, B. Gara, 4 calanoid copepods were adjusted to ranging from N.R. Bromage and J.R. Sargent, to cells mlg dayg [44]. The lower densities in Natural copepods are superior to enriched e present study may also be attributed to e lower food Artemia nauplii as feed for halibut larvae concentrations compared to oer studies. It seems at (Hippoglossus hippoglossus) in terms of survival, fecundity has been decreased in lower food pigmentation and retinal morphology: relation to concentrations [30]. dietary essential fatty acids. Journal of Nutrition, The intensive production of e harpacticoid and 29: cyclopoid copepods were reported in different literatures 3. Hernandez Molejon, O.G. and L. Alvarez-Lajonchere, and resulted wi a higher density production compared Culture experiments wi Oiona aculata to calanoid copepods. Harpacticoids such as species of Farran, 93 (Copepoda: Cyclopoida) and its genus of Nitokra, Tisbe, Euterpina, Tigriopus and advantages as food for marine fish larvae. Amphiascoides are e preferred organisms for e Aquaculture, 29: development of an intensive copepod culture system 4. Van der Meeren, T., R.E. Olsen, K. Hamre and [,6,8,7,45]. For example, [45] harvested a daily average H.J. Fyhn, Biochemical composition of 2 yield of 300,000 nauplii per tray or 25 nauplii cmg dg for copepods for evaluation of feed quality in harpacticoid copepod, Tisbe spp. [7] also reported production of juvenile marine fish. Aquaculture, 2 harvests of 70 copepods cmg dg from a marine 274: harpacticoid copepod in mass culture of Amphiascoides 5. Toledo, J.D., M.S. Golez, M. Doi and A. Ohno, 999. atopus in a recirculating system culture. In general, Use of copepod nauplii during early feeding harpacticoids can be grown in densities up to 5,000 stage of grouper, Epinephelus coioides. Fish. Sci., individuals LG [, 46]. 65(3): Støttrup, J., The elusive copepods: CONCLUSION eir production and suitability in marine From e results of is study, it can be concluded 7. aquaculture. Aquaculture Research, 3: Kleppel, G.S., S.E. Hazzard and C.A. Burkart, at A. clausi could survive at a very broad range of Maximizing e nutritional values of copepods in salinities (3-45 as achieved in e current study). This aquaculture: Managed versus balanced nutrition. study was conducted at laboratory scale over relatively In: Copepods in Aquaculture, Eds., C.S. Lee, short periods of time; erefore e results may not be P.J. O'Bryen and N.H. Marcus, Blackwell Publishing, fully reproducible in large-scale cultures wi capacity of Ames, Iowa, pp: m tanks. However, it clearly served e purpose of 8. Drillet, G., S. Frouël, M.H. Sichlau, P.M. Jepsen, identifying e optimum salinity conditions for culturing J.K. Højgaard, A.K. Joarder and B.W. Hansen, 20. of A. clausi, which are likely to be applicable in larger- Status and recommendations on marine copepod scale cultures. Based on e findings of is study, to cultivation for use as live feed: A review. achieve maximum density of A. clausi culture for Aquaculture, 35: aquaculture purposes, it could be kept and cultured at a 9. Payne, M.F., R.J. Rippingale and R.B. Longmore, salinity of Grow and survival of juvenile pipefish, Stigmatopora argus fed live copepods wi high and ACKNOWLEDGEMENT low HUFA content. Aquaculture, 67: Doi, M., J.D. Toledo, M.S.N. Golez, M. Santos and The research assistance provided by SANRU A. Ohno, 997. Preliminary investigations of feeding University. performance of larvae of early red-spotted grouper, Epinephelus coioids reared wi mixed zooplankton. REFERENCES Hydrobiologia, 358: Payne, M.F. and R.J. Rippingale, 2000a. Rearing West. Lee, C.S., P. O Bryen and N.H. Marcus, Australian seahorse, Hippocampus subelongatus Copepods in Aquaculture. Blackwell Publishing, juveniles on copepod nauplii and enriched Artemia. pp: 352. Aquaculture, 88:

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