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Aquaculture 232 (2004) 637 – 649 www.elsevier.com/locate/aqua-online Evaluation of egg production and quality in the
Aquaculture 232 (2004) 637 – 649 www.elsevier.com/locate/aqua-online Evaluation of egg production and quality in the

Aquaculture 232 (2004) 637 – 649

Aquaculture 232 (2004) 637 – 649 www.elsevier.com/locate/aqua-online Evaluation of egg production and quality in the

www.elsevier.com/locate/aqua-online

Evaluation of egg production and quality in the Mediterranean red porgy (Pagrus pagrus ) during two consecutive spawning seasons

Constantinos C. Mylonas a, * , Maria Papadaki a,b , Michalis Pavlidis b , Pascal Divanach a

a Institute of Marine Biology of Crete, P.O. Box 2214, Iraklion, Crete 71003, Greece b Department of Biology, University of Crete, P.O. Box 2208, Iraklion, Crete 71409, Greece

Received 5 December 2002; received in revised form 11 July 2003; accepted 14 July 2003

Abstract

Egg production from two broodstocks of captive-reared red porgy ( Pagrus pagrus) was monitored for two consecutive seasons (2001 and 2002). Spawning lasted from January to April –May under water temperature between 15.8 and 19.2 jC. Spawning was mostly daily, though over the course of the two spawning seasons, the mean interval was 1.5 – 1.2 days spawn 1 . There was a large variation in daily fecundity without a consistent trend, except for a slight peak in the middle of the spawning season. Mean (S.E.M.) annual relative fecundity ranged between 408,300 (95,800) and 442,500 (180,800) eggs kg 1 year 1 . Mean fertilization success increased significantly from 36.9 (2.0)% in 2001 to 68.9 (2.5)% in 2002, resulting in mean annual viable egg production of 174,800 (87,600) and 301,500 (99,800) eggs kg 1 , respectively. There were no significant differences between the 2 years in other egg quality parameters, which ranged between 74.5 and 79.3% 1-day embryo survival, 81.6 – 85.5% hatching success and 76.7 – 78.8% 5-day larval survival. Of the relations examined, fertilization % had a significantly positive correlation with 1-day embryo survival, which in turn was significantly correlated with hatching %. Larval survival was further correlated with hatching %. These data suggest that decisions on whether to proceed with the incubation of a batch of eggs and rearing of the larvae can be made within 1 day from egg collection, before much effort is invested by the hatchery. The present study presents further information on the spawning kinetics, and egg production and quality characteristics of the Mediterranean red porgy in culture, and demonstrates the suitability of this species for reproduction in captivity. D 2004 Elsevier B.V. All rights reserved.

Keywords: Pagrus; Red porgy; Spawning; Egg quality

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1. Introduction

The red porgy ( Pagrus pagrus L. 1758) is a demersal marine fish widely distributed in the Mediterranean Sea and Atlantic Ocean, and of great commercial importance to the fishing industry (Manooch and Hassler, 1978; Vassilopoulou and Papaconstantinou, 1992; Harris and McGovern, 1997). It is considered a species of great commercial importance for the aquaculture industry, due to its wide geographical distribution, high market demand and good growth rates (Manooch and Hassler, 1978; Kentouri et al., 1995; Pajuelo and Lorenzo, 1996; Maragoudaki et al., 1999) . As a result, in the past decade, many commercial hatcheries begun experimenting with its broodstock management, larval rearing and grow out (Kolios et al., 1997; Abellan and Basurco, 1999; Bodington, 2000; Mihelakakis et al., 2001) . A close relative of the red porgy, the red seabream ( Pagrus major) has been cultured successfully for many years in Japan, where it is the most important cultured species of the Sparidae family (Foscarini, 1988; Watanabe and Kiron, 1995) . Also, efforts are underway for the study and establishment of another congener, the snapper ( Pagrus auratus ) in New Zealand and Australia (Scott et al., 1993; Pankhurst, 1994; Cleary et al., 2002). Despite similar efforts with the red porgy, commercial production has not yet reached the scale of other marine fishes in the Mediterranean, due to problems with its skin discoloration in grow out facilities (Kolios et al., 1997; Cejas et al., 2003; Pavlidis et al., 2002) . The red porgy is a protogynous diandric hermaphroditic species, with all fish possessing an ovary or ovotestis with a dominant ovarian territory during the first 2 years of life (Kokokiris et al., 1999; Fostier et al., 2000) . Puberty in captivity, both in females and primary males, is first observed in 3-year-old fish, but even at 4 years of age, only 50% of females may be reproductively mature (Kokokiris et al., 1999) . The various stages of gametogenesis during the annual reproductive season, along with the associated plasma vitellogenin (Kokokiris et al., 2001) and sex steroid hormones, have been recently described (Kokokiris et al., 2000). Red porgy display an asynchronous ovarian develop- ment and spawn, depending on latitude, from February to mid-June (Manooch, 1976; Vassilopoulou and Papaconstantinou, 1992; Vaughan et al., 1992; Pajuelo and Lorenzo, 1996; Mihelakakis et al., 2001) . However, precise data on spawning kinetics, fecundity and egg quality in captivity are still lacking. The observed varying egg and larval quality produced in fish farms is considered an important limiting factor on fry production (Kjørsvik et al., 1990) and, therefore, the further development of aquaculture (Bromage, 1995) . Egg quality from the aquaculturist’s point of view can be defined as the egg’s potential to produce a viable fry (Kjørsvik et al., 1990) . Egg quality can be influenced by various parameters that often change during the spawning season. These may include endocrine status of the female during oogenesis, feed ration and quality, physico-chemical parameters of the rearing water, broodstock handling stress, etc. (Campbell et al., 1992; Bromage, 1995; Brooks et al., 1997; Christiansen and Torrissen, 1997; Carrillo et al., 2000) . Therefore, it is important for every species with a potential for aquaculture to investigate the existence of seasonal as well as inter-annual variations in egg quality, in order to optimise egg collection and larval production in commercial hatcheries. The objectives of the present study were to obtain further information on the spawning kinetics, egg production and quality characteristics of red porgy over two complete

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spawning seasons in captivity. In addition, the existence of a relation between various egg production, embryo and larval survival parameters was examined, in order to identify potential predictors of larval survival.

2. Materials and methods

2.1. Broodstock and husbandry

Two groups of wild-caught red porgy (4 –5 years old) were maintained in 5-m 3 tanks and were exposed to a simulated natural photoperiod and ambient water temperature (16 – 26 j C) beginning in the winter of 2000. A flow-through supply of filtered surface and shallow-well seawater provided a water exchange of >50% h 1 . Fish were fed by means of a self-feeder with commercial extruded feed (Biomar Hellenique S.A., No 8) and three times a week were offered frozen squid. Each tank was fitted with an overflow egg collector, and spontaneously spawned eggs were collected daily during two consecutive spawning seasons (2001 and 2002). In 2001, the first group (R1a) contained eight females and three males and the second group (R1b) contained five females and three males of mean (S.D.) weight of 3.07 (0.5) kg and 2.89 (0.4) kg, respectively. In 2002, after some mortality during the previous year, group R1a contained six females and four males (one female sex-inverted), and group R1b contained four females and three males of mean weight of 3.0 (0.3) kg and 3.4 (0.8) kg, respectively. The mean ambient water temperature during the spawning period during both years was 16.7 (0.6) j C, ranging between 15.8 and 19.2 j C.

2.2. Evaluation of egg and larval quality

Eggs were obtained from the egg collectors and were initially placed in a 10-l bucket in order to estimate fecundity and fertilization % after sub-sampling of 10 ml. On a daily basis during the 2001 spawning season, oocyte diameter and oil globule diameter of 12 eggs from each tank were measured with the use of a microscope fitted with an ocular micrometer. In order to monitor embryo and larval survival, eggs from two to three spawns every week were individually placed in 96-well microtiter plates (in replicates) according to the procedure of Panini et al. (2001) , with some modifications. Briefly, floating (>90% fertilized) eggs from the 10-l bucket were taken in a 250-A m mesh filter and were rinsed with sterilized seawater and poured in a 2-l beaker. A Petri dish was used to scoop 100 – 200 eggs from the beaker. The Petri dish was then placed under a stereoscope and only fertilized eggs were taken one by one with a micropipette set to 200 A l and transferred to the wells of the microtiter plates (one egg per well). The microtiter plates were then covered with a plastic lid, placed in a controlled-temperature incubator and maintained for 5 days at 16 ( F 0.5) j C. Using a stereoscope, embryonic and early larval development was evaluated once a day, recording the number of 1-day embryos, hatched larvae and viable larvae on day 5 after egg collection. For analysis, various egg quality parameters were evaluated. Fertilization success was evaluated daily as soon as the eggs were obtained from the egg collectors and was

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calculated as the number of fertilized eggs/total spawned eggs. Embryo survival was evaluated the day after egg collection (1 day), and was calculated as the number of eggs having live embryos/the number of fertilized eggs initially loaded in the microtiter plates. Hatching success was calculated as the number of hatched larvae/the number of live 1-day

as the number of hatched larvae/the number of live 1-day Fig. 1. Daily relative fecundity (eggs
as the number of hatched larvae/the number of live 1-day Fig. 1. Daily relative fecundity (eggs

Fig. 1. Daily relative fecundity (eggs kg 1 female biomass) and fertilization success of two red porgy broodstocks (R1a and R1b) during the spawning season of 2001. The data shown in the figure represent all the spawns observed during the reproductive period. Fish were maintained in 5-m 3 tanks supplied with filtered water of a mean (S.D.) temperature during the spawning season of 16.7 (0.6) jC. The R1a and R1b broodstocks consisted of eight females: three males and five females: three males, and the total female biomass was 22.7 and 14.1 kg, respectively.

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C.C. Mylonas et al. / Aquaculture 232 (2004) 637–649 641 Fig. 2. Daily relative fecundity (eggs
C.C. Mylonas et al. / Aquaculture 232 (2004) 637–649 641 Fig. 2. Daily relative fecundity (eggs
C.C. Mylonas et al. / Aquaculture 232 (2004) 637–649 641 Fig. 2. Daily relative fecundity (eggs

Fig. 2. Daily relative fecundity (eggs kg 1 female biomass) and fertilization success of two red porgy broodstocks (R1a and R1b) during the spawning season of 2002. The data shown in the figure represent all the spawns observed during the reproductive period. Fish were maintained in 5-m 3 tanks supplied with filtered water of a mean (S.D.) temperature during the spawning season of 16.7 (0.6) jC. The R1a and R1b broodstocks consisted of six females: four males and four females: three males, and the total female biomass was 20.2 and 12.7 kg, respectively.

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embryos, and 5-day larval survival was calculated as the number of live larvae 5-day after egg collection/the number of hatched larvae. Estimating survival % by using in the denominator, the number of individuals that survived to the previous developmental stage (Mylonas et al., 1992) results in a more independent evaluation of survival within specific developmental stages, without the potential of a masking effect of the previous stage.

2.3. Statistical analysis

Differences in spawning and egg quality between the two spawning seasons were examined using one-way Analysis of Variance (ANOVA) at a minimum significance of P < 0.05. The existence of a correlation between some egg quality parameters was examined using Simple Regression analysis. Unless otherwise mentioned, results are presented as means (S.E.M.). Statistical analyses were done with a linear statistics software (SuperAnova, Abacus Concepts, USA).

3. Results

3.1. Spawning and egg quality

The spawning season of red porgy lasted from 14 January to 14 May in 2001 (Fig. 1) and from 19 January to 18 April in 2002 (Fig. 2) . There was a large variation in daily relative fecundity without a consistent trend through the spawning season. Egg fecundity in tank R1a showed a slight increase towards the middle of the spawning season during both years (Fig. 1) . Similar increases in egg fecundity were observed also in tank R1b, but the 2002 spawning period of this stock was shorter and occurred towards the last part of the season (Fig. 2) . Spawning interval ranged from 1 to 9 days, but the vast majority of spawns occurred within 1 –2 days from each other, resulting in a mean spawning interval of 1.5 (0.2) days for 2001 and 1.2 (0.1) days for 2002 (data not shown). Fertilization success exhibited larger daily variation in 2001 (Fig. 1) compared to 2002 (Fig. 2) , and mean fertilization success differed significantly ( P = 0.001) between the 2 years, increas- ing from 36.9% in 2001, to 68.9% in 2002 (Table 1) .

Table 1 Cumulative spawning parameters of red porgy broodstock (n = 2) from two consecutive spawning seasons (2001 – 2002)

Mean (S.E.M.)

2001

2002

Fecundity (1000 eggs kg 1 ) Fertilization % Viable egg production (1000 eggs kg 1 ) a 1-day embryo survival % Hatching % 5-day larval survival %

408.3 (95.8)

442.5 (180.8)

36.9 (2.0)

68.9 (2.5)

174.8 (87.6)

301.5 (99.8)

79.3 (8.4)

74.5 (2.7)

81.6 (13.4)

85.5 (11.6)

78.8 (2.0)

76.7 (5.1)

There were no statistically significant differences between the 2 years in the parameters examined (ANOVA, P < 0.05), except for fertilization % ( P = 0.001). a Total number of fertilized eggs produced.

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C.C. Mylonas et al. / Aquaculture 232 (2004) 637–649 643 Fig. 3. Linear correlation between various
C.C. Mylonas et al. / Aquaculture 232 (2004) 637–649 643 Fig. 3. Linear correlation between various
C.C. Mylonas et al. / Aquaculture 232 (2004) 637–649 643 Fig. 3. Linear correlation between various
C.C. Mylonas et al. / Aquaculture 232 (2004) 637–649 643 Fig. 3. Linear correlation between various

Fig. 3. Linear correlation between various production (relative fecundity, fertilization success) and survival parameters (1-day embryo survival, hatching success and 5-day larval survival) of red porgy eggs collected throughout the spawning seasons of 2001 and 2002. Where there is statistically significant correlation, the significance value ( P) and correlation coefficient ( r 2 ) is shown. Absence of correlation is indicated by ‘‘ns’’ (nonsignificant).

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Mean annual relative fecundity ranged from 408,300 to 442,500 eggs kg 1 female body weight, and did not differ significantly between the two spawning seasons (Table 1) . The resulting mean total fertilized egg production was 174,800 eggs kg 1 for 2001 and 301,500 eggs kg 1 for 2002 (Table 1) . Except for fertilization %, no significant differ- ences between the 2 years were observed on egg quality parameters such as 1-day embryo survival, hatching success or 5-day larval survival (Table 1) . Regression analysis indicated that there was no correlation between fertilization % and fecundity during neither of the two spawning seasons (Fig. 3). On the other hand, there was significant correlation between 1-day embryo survival and fertilization %, and between hatching % and 1-day embryo survival, during both spawning seasons. The high significance obtained in this correlation was mostly due to a few points at the extreme low range. However, even when these points were removed from the analysis, the relation between hatching % and 1-day embryo survival still remained significant with P < 0.003 and P < 0.02 for the 2001 and 2002 seasons, respectively (data not shown). Finally, 5-day larval survival correlated significantly with hatching % in 2001, but not in 2002. There was no consistent trend in changes in oocyte or lipid droplet diameter during the spawning season, which ranged between 1.07 and 0.99 mm, and between 0.24 and 0.22 mm, respectively (data not shown). Over the whole spawning season, mean (S.D.) egg diameter was 1.02 (0.02) mm and mean oil globule diameter was 0.228 (0.004) mm.

4. Discussion

The spawning period of red porgy in the present study span from mid-January to early-May in 2001 (Fig. 1) and from mid-January to mid-April in 2002 (Fig. 2) . Except from one broodstock in 2002 (R1b), in both years the season begun earlier than reported from other cultured or wild populations, which in general exhibit limited temporal variation in spawning season, despite the very wide geographical distribution of this species. For example, in the Atlantic Ocean, the spawning period extends from February to May (Manooch, 1976; Pajuelo and Lorenzo, 1996) , while in the eastern part of the Mediterranean Sea, spawning spans between February and May (Vassilo- poulou and Papaconstantinou, 1992) . Also, reported spawning seasons for captive- reared stocks of Mediterranean red porgy were from mid-February to early June (Mihelakakis et al., 2001) and from mid-February to mid-May (Kolios et al., 1997) . Finally, using another captive broodstock in the same facilities as the present study, Kokokiris et al. (2000) reported spawning activity between March and May, while spermiation begun in February (Kokokiris et al., 2001) . It is unclear why the spawning season occurred earlier in the stocks used in the present study, but it may be the result of differences in the spawning conditions between captivity and the wild. In the wild, reproductively mature fish occupy deep waters (50 –150 m) where the temperature is lower and more stable during the year (14.5 –16.5 j C) (Machias et al., 1998). In the present study, the fish were exposed to surface seawater fluctuations ranging from 15.5 to 24.0 j C. It is possible that cultured fish may undergo rapid maturation and begin to spawn as soon as appropriate spawning water temperatures of below 20 j C (Manooch and Hassler, 1978) appear in early winter. Similarly, spawning of red porgy in the

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Canary Islands occurs at water temperatures between 18 and 20 j C (Pajuelo and Lorenzo, 1996) . Although daily fecundity did not differ significantly between the two broodstocks, during both monitoring years, there were obvious differences in the number of spawns, as well as the timing of the spawning period (Figs. 1 and 2) . The higher number of spawns per year in R1a could be attributed to the greater number of females that constituted this broodstock, compared to R1b. For most of the spawning period, spawning occurred daily, but it is not known if every individual female red porgy spawns every day. Based on the spawning kinetics and large daily fecundity variations exhibited by both broodstocks, at least for part of the season, it appears likely that not all females spawn in a particular day. Therefore, a broodstock constituted by a larger number of females is more likely to produce spawns every day of the spawning season. In terms of the observed differences in the onset and duration of the spawning season among the two broodstocks, we cannot explain the fact that spawning of R1b in 2002 did not start until March and was completed before R1a. The two broodstocks were maintained in exactly the same type of tank, located side-by-side, supplied by the same water source and exposed to the same lighting conditions. Variations in the duration of spawning, spawning peak and fecundity have also been reported among different broodstocks, as well as in the same broodstock between consecutive spawning seasons, in another study of Mediterranean red porgy (Kentouri et al., 1995). Taken together, the above information underlines some problems which may be encountered in managing cultured red porgy broodstocks, namely variations in the spawning kinetics and egg production characteristics among seemingly identical stocks, as well as in the same broodstock between different spawning seasons. In terms of annual relative fecundity, the values obtained here are lower than the 770,000 eggs kg 1 reported by Mihelakakis et al. (2001) , but higher than the 200,000 eggs kg 1 reported in another study (Kolios et al., 1997). In the congener red seabream, annual relative fecundity reportedly ranges between 260,000 eggs kg 1 (Kafuku and Ikenoue, 1983) and 660,000 eggs kg 1 (Watanabe and Kiron, 1995) . Fertilization success showed large variations over the course of the spawning season (Figs. 1 and 2) , especially during the first year of monitoring, exhibiting significant differences in mean values between the two spawning seasons. During 2002, fertilization success increased towards the middle of the spawning season in broodstock R1a and remained high until the end of the season, whereas in broodstock R1b, it remained at high levels throughout the spawning season. Overall, the mean fertilization success in the present study was 36.9% and 68.9% for 2001 and 2002, respectively (Table 1) , which compares well with other studies of cultured red porgy. Kentouri et al. (1995) , monitoring the spawning performance of two broodstocks over 2 consecutive years, reported production of between 2% and 60% viable floating eggs, (of which >95% are usually fertilized). In another study, a large broodstock of red porgy (100 kg female biomass) maintained at low densities in a 42-m 3 outdoor tank produced eggs of mean fertilization success of >80% (Mihelakakis et al., 2001) . In the latter study, an improvement of fertilization % was obtained towards the peak and decreased towards the end of the spawning season, something that was not observed here. The increase in fertilization % and improvement in total number of viable egg production from a mean of 174,800 in 2001 to 301,500 eggs kg 1 in 2002 (Table 1) may suggest a better acclimatization of the

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wild-caught broodstocks to the rearing tanks. It may also be due to the sex ratio, which in 2002 was less skewed in favor of males and more towards a 1:1 compared to 2001, an effect also observed in another sparid fish, the common dentex (Dentex dentex ) (Pavlidis, 2000) . The observed varying egg and larval quality produced in fish farms of various species can be considered as an important limiting factor on the development of aquaculture (Kjørsvik et al., 1990; Bromage, 1995) . The quality of the produced eggs can be influenced by various parameters which can change during the spawning season, such as feed ration and quality, water temperature and quality, handling stress, etc. (Kjørsvik et al., 1990; Campbell et al., 1992; Bromage, 1995; Christiansen and Torrissen, 1997; Carrillo et al., 2000) . It is useful, therefore, to be able to determine the quality of the eggs produced in a particular batch and predict the survival of the larvae, before investing facilities, manpower and time in incubating eggs and rearing larvae. Some of the egg quality markers employed in different fishes include fecundity, buoyancy of pelagic eggs, fertilization success, morphological characteristics (cleavage pattern or distribution of oil globules), eyeing % (in salmonids) and hatching success (Kjørsvik et al., 1990; Bromage and Roberts, 1995; Brooks et al., 1997; Shields et al., 1997; Nocillado et al., 2000) . In the present study, the existence of a correlation between various egg production, embryo and larval survival parameters was examined, in order to identify potential markers of egg quality in red porgy. Of the relations examined during two consecutive spawning seasons in two separate broodstocks, it was found that fertilization % had a statistically significant positive correlation with 1-day survival of the embryos, which in turn was significantly correlated with hatching % (Fig. 3) . Larval survival 5-day after spawning was also correlated with hatching %, but this relation existed only during the 2002 season. Although the coefficients ( r 2 ) for these correlations were low, these findings suggest that larval survival prior to the onset of exogenous feeding can be predicted to a certain extend by the fertilization % or the 1-day embryo survival of the particular batch of eggs. Especially regarding the relation between hatching % and 1-day embryo survival, it is important to note that batches of eggs with >60% 1-day embryo survival always had >80% hatching success. Therefore, decisions on whether to proceed with the incubation of the eggs and rearing of the larvae can be made within 1 day from egg collection, before much effort is invested by the hatchery. Various egg quality predictors have been identified in other cultured fishes. For example, in the European sea bass ( Dicentrarchus labrax ) hatching success and 4-day larval survival exhibited a significant positive correlation with 1-day embryo survival (Mylonas et al., 2003) . The symmetry of cell division at the early blastula stage is considered a strong predictor of hatching and normal larval development in cod ( Gadus morhua) (Kjørsvik, 1994) , halibut ( Hippoglossus hippoglossus ) (Shields et al., 1997) and other marine fishes (Kjørsvik et al., 1990) , including the red seabream (Sakai et al., 1985) . Finally, the easiest to apply and best studied egg quality determinant has been reported in salmonids, where fertilization success correlates very well with subsequent eyeing and hatching %, as well as larval survival (Bromage and Cumarana- tunga, 1988) . Manooch (1976) first reported that red porgy eggs are transparent and spherical in shape, their diameter is 0.8– 0.9 mm and have a centrally located oil globule of 0.19– 0.21 mm. Kolios et al. (1997) found mean egg diameter to be 0.84 mm for fish kept in captivity

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at 13 –25 j C, whereas Mihelakakis et al. (2001) observed egg diameters from 0.99 to 1.09

mm during one spawning season. Egg and lipid droplet diameter of red porgy eggs in the

present study ranged between 1.07 and 0.99 mm, and between 0.24 and 0.22 mm, respectively (data not shown). The different egg dimensions found are not contradictory, considering that broodstock size and genotype, as well as the daily and seasonal rates of food provision can influence the diameter of produced eggs (Bromage, 1995) . In the red seabream, normal eggs can range in diameter between 0.66 and 1.03 mm, without any reported effect on egg quality (Watanabe and Kiron, 1995) . Unlike an earlier published report on red porgy (Mihelakakis et al., 2001) , significant changes in oocyte diameter were not observed in the present study. A reduction in egg diameter, and therefore yolk content, may occur towards the end of the spawning period in fishes with asynchronous ovarian

development (Mihelakakis et al., 1995, 2001) , and may result in a reduction in the survival % of the embryo and larvae (Blaxter, 1988) . No such reduction was observed in the present study, since 1-day survival, hatching success and 5-day larval survival did not show any consistent trend during the spawning period, and averaged above 74%, 80% and 75%, respectively, during the 2 years of the study (Table 1) . The above results demonstrate

that one can expect eggs of similar quality to be produced at any time during the spawning

season of red porgy.

The present study presents new information on the spawning kinetics, egg production

and quality characteristics of the Mediterranean red porgy in culture, and provides further

evidence for the suitability of this species for reproduction in captivity. Such information is valuable for the proper organization of a commercial hatchery, specifically as it regards

broodstock biomass requirements, egg production schedules, and predictors of egg quality

and larval survival. Further research on the reproduction of red porgy in captivity should

focus on the effect of broodstock age, sex ratio and nutrition on egg production and

quality.

Acknowledgements

This work was carried out during a University of Crete, post-graduate laboratory exercise of M.P. at the Department of Aquaculture, Institute of Marine Biology of Crete, under the supervision of Professor Maroudio Kentouri. We would like to thank Irini Sigelaki for the broodstock husbandry and sampling assistance.

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