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Effects of Photoperiod on Growth of Larvae and Juveniles of the Anemonefish melanopus M. Arvedlund, M.I. McCormick and T. Ainsworth

Abstract Rearing of anemonefishes is now relatively routine compared to the culture of cardinalfishes () or angelfishes (Pomacanthidae). However, it is still a labor intensive, time intensive and expensive procedure. To reduce time and cost of rearing anemonefishes, experiments were undertaken to improve the methods for rearing Amphiprion melanopus. These experiments were conducted to determine the effect of the length of photoperiod on larval duration, growth to metamorphosis and early juvenile phase. Growth of larvae was significantly faster and the duration of the larval phase was significantly shorter, under a photoperiod of 16 hours light/8 hours dark, compared to the photoperiods of 12 hours light/12 hours dark and 24 hours light/0 hours dark.

Introduction This paper describes the growth a 16-hour daily light period, while experiments conducted at the Sir Juhl (pers. comm.) recommends a 24- The tropical marine anemone- George Fisher Research , hour light regime. None of these au- () are important James Cook University (JCU), to im- thors present supporting data. This in the trade for ornamental prove the methods for rearing study investigates the effect of 12 (Wilkerson 1998) and are a popular anemonefishes so as to reduce the hour, 16 hour and 24 hour photo- subject of research (Fautin 1991). time and cost of rearing them for sci- periods on the growth and larval Over the last 20 years, mariculture entific studies. duration of the anemonefish A. centers and scientific laboratories Only two studies have experi- melanopus. have started rearing these fishes in mented with ways of enhancing the large quantities (McLarney 1985, efficiency of larval rearing of 1986; Miyagawa 1989; Hoff 1993; anemonefishes. Frakes and Hoff Materials and Young 1996; Job et al. 1997). The (1982) published a study on the ef- Methods list of marine fishes reared in cap- fect of high nitrate-N on the growth FISH MAINTENANCE tivity today, for purposes other than and survival of juvenile and larval AND REARING human consumption, contains more anemonefish A. ocellaris. Alayse than 84 (Tables 1 and 2). The (1984) studied the survival rate of Breeding pairs of A. melanopus fact that 26 different species from the A. ocellaris larvae fed on enriched were collected from the Cairns sec- family Pomacentridae are reported to food. This study examines the tion of the and be reared is notable. This is a sig- growth rates of larvae and juveniles placed in 60-l tanks with gravel fil- nificantly higher number of species under different light regimes. This ters and 200-l powerheads for the compared to all other families. How- variable is important as anemonefish circulation of the water. Rocks were ever, when we look at species that larvae are visual feeders (Coughlin placed as a surface on which the fish can be reared reliably in large quan- 1994; Job and Bellwood 1996). Sci- could . Water in the tanks was tities, they include only a dozen entists have shown that an extended obtained from the JCU aquarium sys- anemonefish species, seven species photoperiod can significantly in- tem. It had 33% salinity, 27-30°C tem- of gobiids (Gobiidae), five species of crease the growth rate of the larvae perature (daily variation, summer), cardinalfishes (Apogonidae) and and early juveniles of a variety of 21-25°C (daily variation, winter) and eight species of pseudochromids marine fish species (Fuchs 1978; a pH of 8.0-8.2. The water in the pa- (Pseudochromidae) The last two are Barahona-Fernandes 1979; Boehlert rental tank was flushed daily with only included here as a result of re- 1981; Kiyonon and Hirano 1981; water from the main aquarium sys- cent work by Gardner (1997) and Job Tandler and Helps 1985; Duray and tem. The JCU system is closed, with et al. (1997). For all the species Kohno 1988; Barlow et al. 1995). coastal water filtered through sand within the families mentioned Hoff (1996) and Wilkerson (1998) and large protein skimmers. above, larval rearing is still time-con- estimate the optimum growing con- Spawning occurred approximately suming and expensive. ditions for anemonefish larvae to be every three weeks, and produced

18 Naga, The ICLARM Quarterly (Vol. 23, No. 2) April-June 2000 Naga, TheICLARM Quarterly(Vol. 23,No.2)April-June2000 Labridae Tetraodontidae (8 species) Syngnathidae (3 species) Serranidae (4 species) Sciaenidae (9 species) Pseudochromidae (2 species) Pomadasyidae Pomacentridae (6 species) Pomacanthidae Plesiopidae Opistognathidae (2 species) Lutjanidae (10 species) Gobiidae Gobiesocidae Ephippidae Callionymidae Carangidae Blennidae Batrachoididae (6 species) Apogonidae Table 1. List of marine fishes reared in captivity for purposes other than human consumption. human than other purposes for captivity in reared fishes marine of List 1. Table aiySeisCommonname Species Family Lachnolaimus maximus Spoeroides maculatus Syngnathoides biaculatus Hippocampus zostera Hippocampus reidi Hippocampus punctulatus Hippocampus kuda Hippocampus hippocampus Hippocampus erectus Doryrhampus dactyliophorus Hypoplectrus unicolor Gramma melacara Gramma loreto Equetus umbrosus Equetus punctatus Equetus lanceolatus Equetus acuminatus Pseudochromis sankey Pseudochromis olivaeceous Pseudochromis flavivertex Pseudochromis aldabrensis Ogilbyina novaehollandiae Labracinus cyclophtalmus Haemulon plumieri Anisotremus virginicus (26 species)SeeTable 2. Pomachantus paru Pomachantus arcuatus Centropyge resplendens Centropyge loriculus Centropyge ferrugatus Centropyge argi Calloplesiops altivelis Opistognathus aurifrons Ocyurus chrysurus Lutjanus griseus Gobiosoma okinawae Coryphopterus personatus Gobiosoma genie Gobiosoma prochilus Gobiodon citrinus Elactinus xanthipora Gobiosoma oceanops Pterapogon kauderni Apogonichtys nigripinnis Gobiosoma evelynae Gobiosoma multifasciatum Gobiesox strumosus Chaetodipterus faber Synchiropus spendidus Trachinotus carolinus Blennius pavo Opsanus tau Cheilodipterus quinquelineatus Sphaeramia nematoptera Apogon compressus Apogon cyanosoma ofs Moe1992 Moe1992 Hogfish Northern pufferfish Pipefish pte ehreLange1989 Moe1992 Dwarf seahorse seahorse Spotted seahorse Lined Ringed pipefish altMoe1992 Hamlet rmAnon.1997 Moe1992 Moe1992 Moe1992 Blackcap basslet Royal gramma Drum Spotted drum Jacknife-fish High-hat Springeri Striped dottyback Olive dottyback Yellow dottyback Orchid dottyback Sunrise dottyback Neon dottyback ht rn Moe1992 Australian dottyback Dottyback White grunt Porkfish ryaglihMoe1975 French angelfish angelfish Grey oe-aiebtaWassink1990 Y Flame angel Comet-marine betta Yellowhead jawfish Yellowtail snapper Grey snapper odngb Y Anon. 1997 Yellow goby Masked goby Genie’s cleaninggoby West indiancleanergoby Citron goby Golden goby Neon goby Marini1996 cardinal Banggai Nigripes cardinal klefs Moe1992 Walker1991 Schumann1969 Sharknosed goby Patzner Greenband goby Skilletfish spadefish Atlantic Job(pers.comm.) Mandarinfish Florida pompano Jobetal.1997 Mediterranean blennnius Jobetal.1997 toadfish Common Fiveline-cardinal Pyjama cardinal Split-banded cardinal Yellow-stripedcardinal Lange 1989 Moe 1992 Lange 1989 Anon. 1998 Schumann 1969 Lange 1989 Moe 1992 Moe 1992 Gardner 1997 Gardner 1997 Gardner 1997 Gardner 1997 Brons 1996 Brons 1996 Gardner 1997 Gardner 1997 Lange 1989 Moe 1992 Moe 1975 Anon. 1998 Anon. 1998 Moe 1992 Moe 1992 Hioki etal.1990 Anon. 1997 Gardner 1997 Anon. 1997 Anon. 1997 Anon. 1997 Moe 1992 Lange 1989 Moe 1992 Moe 1992 Gardner 1997 Moe 1992 and Brandstaetter1989 Job etal.1997 oung 1982 oung 1994 Reference 19

AQUABYTE 200 - 300 eggs per clutch. Embryos seen. These methods of reducing chopped sardines, prawns and vi- hatched in nine days. An hour or light stopped the “headbutting syn- tamin supplements, i.e., the same two before hatching, the rock with drome” of the fish and improved diet as adult fish. the egg clutch (and the sea water quality, since the act as anemone) was transferred in a wa- a nutrient sink (Job et al. 1997). GROWTH AND SURVIVAL ter-filled bucket to the hatching Water in the hatching tank came from aquarium, where the clutch was left the breeding tank to ensure constant Due to the difficulty of dividing in the dark for approximately 90 osmolarity for the fish larvae. Ap- up a batch, larvae from three different min. The water in the hatching tank proximately 20% of the water was batches from the same parents were was gently aerated but not filtered, replaced every second day with used for the experiments. To examine since the fish larvae are sensitive to water from the parent aquarium. variability in growth among different currents (Arvedlund, pers. obs.). The larvae were fed the rotifer batches of larvae from a single pair of The sides of the hatching tanks were Brachionus plicatilis for the first two A. melanopus, three batches of larvae covered with black plastic to reduce days after hatching. Then they were from the same parents were exposed light reflection. The phytoplankter gradually introduced to a diet of to a uniform rearing environment Nannochloropsis sp. was used to Artemia. After approximately 30 (16L:8D). These were the control “green up” the tanks until the bot- days, the juveniles were gradually batches. Five fish were randomly tom of the tank could no longer be weaned to a mixed diet of finely sampled every fifth day, starting from the day of hatching (day 0). The sample fish were anesthe- tized by refrigerator chilling, pre- Table 2. List of marine fishes of the family Pomacentridae reared in captivity. served in 70% ethanol and measured to the nearest 0.01 mm total length Species Common name Reference (TL). For wet and dry weight, all fish 1) Anemonefishes (subfamily ) were weighed to the nearest 0.0001 Skunk anemonefish Moe 1992 g. For dry weight, the larvae were Amphiprion allardi Allard’s anemonefish Terver 1975 dried in an oven at 60°C for 16 hours. Barrier Reef anemonefish Fisher (pers. comm.) To examine the effect of extended Amphiprion bicinctus anemonefish Young 1990 photoperiods on fish growth, three Orange-fin anemonefish Moe 1992 Amphiprion clarkii Clark’s anemonefish Miyagawa 1989; Moe 1992; batches of larvae from the same Wilkerson 1992 breeding pair were reared under

Amphiprion ephippium Red saddleback anemonefish Moe 1992; Gardner 1997 three different light regimes. All

Amphiprion frenatus Tomato anemonefish Miyagawa 1989; Juhl 1992; other rearing conditions were repli- Moe 1992; Gardner 1997 cated for each batch. The light re- Wide-band anemonefish Moe 1992 White-bonnet anemonefish Moe 1992 gimes were: (a) 12 hours light/12 hours dark (12L:12D); (b) 16 hours Amphiprion melanopus Red and black anemonefish Moe 1992; Gardner 1997; Job et al. 1997 light/8 hours dark (16L:8D); and (c)

Amphiprion ocellaris False clown anemonefish Miyagawa 1989; Juhl 1992; 24 hours light/0 hours dark Moe 1992; Arvedlund and (24L:0D). The light was provided by Nielsen 1996; Gardner 1997 two 40 watts fluorescent light bulbs. Amphiprion percula Clown anemonefish Moe 1992; Gardner 1997; Job et al. 1997 Food was made available for 24

Amphiprion perideraion Pink anemonefish Miyagawa 1989; Moe 1992; hours/day in all three treatments, in Gardner 1997 densities of 3-5 rotifers (later Amphiprion polymnus Saddleback anemonefish Terver 1971; Moe 1992 Artemia) per ml. A sample of five Amphiprion rubrocinctus Australian anemonefish Moe 1992 Amphiprion sandaracinos Orange anemonefish Miyagawa 1989; fish larvae was collected randomly Moe 1992; Gardner 1997 at hatching (day 0) from each batch, Amphiprion tricinctus Three-band anemonefish Moe 1992 and then every 5 days after hatch-

Premnas biaculatus Spine-cheek anemonefish Moe 1992; Gardner 1997; ing up to day 25. The sample fish Job et al. 1997 were preserved, weighed and mea- 2) Damselfishes other than anemonefishes sured as described above. The test Abudefduf saxitilis Sargeant major Moe 1992 batches were not replicated. Dascyllus albisella Hawaiian Dascyllus Danilowicz and Brown 1992 Growth curves were calculated Dascyllus aruanus Humbug Dascyllus Danilowicz and Brown 1992 for each photoperiod treatment. The Hypsypops rubicundus Garibaldi Moe 1992 growth curves were compared by Microspathodon chrysurus Jewelfish Moe 1992 Neopomacentrus bankieri Chinese demoiselle Job et al. 1997 first using a test for homogeneity of Pomacentrus amboinensis Ambon damselfish Job et al. 1997 slopes. If found nonsignificant, an ANOVA test for differences among

20 Naga, The ICLARM Quarterly (Vol. 23, No. 2) April-June 2000 Naga, TheICLARM Quarterly(Vol. 23,No.2)April-June2000 Tablelowest growth (Fig.2b, 4). with the12L:12Dfishdisplaying the followed bythe24L:0Dtreatment, under 16L:8Dhadthehighest growth differed fromeach other. Fishreared case ofdrybodyweights,all curves ments (F also differedamongphotoperiodtreat- age. Ratesofchangeinbodyweight as changeindrybodyweightwith tuated whengrowthwasexpressed ment (Fig.2a,Table 3). higher slopesthanthe12L:12Dtreat- other, butbothhadsignificantly ments didnotdifferfromonean- from the16L:8Dand24L:0Dtreat- among theslopesfoundthatfish p<0.0008). Pairedcomparisons neity ofslopes:F among treatments(testofhomoge- in totallengthdifferedsignificantly der differentphotoperiods,growth toperiods waslow(Figs.1aand1b). ents butexposedtodifferentpho- batches oflarvaefromthesamepar- used asameasureoflarvalduration. the fish(100%)wasrecordedand hatching andthesettlementofall obs.). Thenumberofdaysbetween (Miyagawa 1989;Arvedlund,pers. midwater, tothebottom ofthetank shift fromoccupyingthetopto development ofwhitebarsanda ogy (Allen1975).Thisinvolvesthe on juvenilebehaviorandmorphol- anemonefish metamorphoseandtake LARVAL DURATION the endofexperiment(day25). batch atthetimeofhatchingand counting thefishlarvaeineach rate inallbatcheswasmadeby tween dryweightandagewaslinear. to analysis.Therelationshipbe- by naturallogtransformationprior was curvilinearandlinearized ship betweentotallengthandage intercepts wasmade.Therelation- These trendswerefurtheraccen- For thethreebatchesrearedun- Variability ingrowthamong At theendoftheirlarvalstage, A grossestimateofthesurvival 2,82 =10.84, p<0.0001).Inthe Results 2,84 =7.71, LARVAL DURATION larvae rearedin16L:8Dalsosettled hatching. Forthetestbatches, gained whitebarsatday8after settled, i.e.metamorphosed,and fish larvae(100%)ofeachbatch mately 70%forallthreebatches. Fig. 1. Regression lines of log transformed total length (a) and dry weight (b) weight dry and (a) length total transformed log of lines Regression 1. Fig. against time for each of three batches of of batches three of each for time against reared under the same conditions. R conditions. same the under reared Lightregime Table 3. Comparison of growth of Comparison 3. Table (total length) versus age for three for light age versus different three length) under (total reared 4:Dy=4.7137Ln(x)+4.5696,R y=4.8933Ln(x)+4.6543, R 24L:0D y=4.0823Ln(x)+4.6486, R 16L:8D 12L:12D batches of of batches regimes. For thethreecontrolbatches,all The survivalratewasapproxi- a b Dry weight (g) Total length (mm) .002 0. 004 0. 006 0. 008 0. A. melanopus .01 0. 10 12 14 4 6 8 0 a a a 0Dy1 a 0Day25 Day 20 Day 15 Day 10 Day 5 Day 0 a 0Dy a1 a1 a2 Day25 Day20 Day15 Day10 Day5 Day0 Equation 000000 000 000 000 000 000 000 000 000 000 000 0000 000 0000 0000 0000 0000 000000 0 0 0 0 0 0 000 000 000 000 000 000000 000000 000000 000000 000000 000000 000000 000000 000000 000000 000000 000000 larvae 2 2 2 =0.9952 =0.9913 =0.9605 2 ³ ³ ³ ³ ³ 0.095 for all equations. all for 0.095 A. melanopus(fromthe16L:8Dgroup) of 16-hourlight.The24-hour est underanextendedphotoperiod up to25daysafterhatchinggrewfast- ter hatching. quired thewhitebarsonday10af- reared in12L:12Dand24L:0Dac- on day8afterhatching,whilethose Lightregime Table 4. Comparison of growth of Comparison 4. Table reared under three different light different three under reared (dry (dry 4:Ly=0.0017x-0.0019, R y=0.002x-0.0023,R 24D:0L y=0.0013x-0.002,R 16D:8L 12D:12L regimes. batches of of batches A. melanopus weight) versus age for three for age versus weight) Log. (16-2) Log. (16-1) Log. (16-3) Discussion Linear (16-2) Linear (16-1) Linear (16-3) A. melanopus larvaeandjuveniles Equation 2 2 larvae 2 =0.9655 =0.9813 =0.9425 light 21 ,

AQUABYTE (Sparus aurata) and Duray and Kohno (1988) for rabbitfish (Siganus

14 guttatus). a The wide range of results sug- gests that different families of fishes 12 have different feeding patterns and therefore different requirements as 10 larvae. Differences in the quality of the food between each study may 8 also play a role. Log. (12 hour) The conclusion of this study is Log. (16 hour) 6 that the optimum lighting condition Log. (24 hour) for growth of A. melanopus is Total length (mm) Total

4 16L:8D, given that appropriate food Day 0 Day 5 Day 10 Day 15 Day 20 Day 25 is present for all the 16 hours of light. Using this photoperiod for lar- 0. 01 val rearing should improve growth rates and decrease larval production

0. 008 time. b

0. 006 Acknowledgement This study is part of Michael 0. 004 Arvedlund’s Ph.D. dissertation con- ducted at James Cook University and

Linear (12 hour) was made possible by doctoral grant 0. 002

Dry weight (g) Linear (16 hour) #1995-154-0080 from the Danish Linear (24 hour) Research Academy, Aarhus, Den- 0 mark. Day 0 Day 5 Day 10 Day 15 Day 20 Day 25 References Fig. 2. Regression lines of log transformed total length (a) and dry Alayse, J.P. 1984. Application of tech- weight (b) against time for each of three batches of A. melanopus, reared under three different light regimes. R2³.095 for all equations. niques used for temperate marine fish in breeding Amphiprion ocellaris (Cuvier). Proc. Mar. Aquar. Oceanogr. Inst. 10:505-519. Allen, G.R. 1975. The anemonefishes. regime yielded faster growth rates light period (Boehlert 1981). A study Their classification and biology. 2nd than the 12L:12D regime. However, on the sea bass Dicentrarchus labrax ed. Tropical Fish Hobbyist, New Jer- both were slower than fish in the reported an optimum growth rate sey. 352 p. 16L:8D photoperiod. The authors with 18-hour light periods Anon. 1997. Breeders register http:// suggest that fish in the extended (Barahona-Fernandes 1979). Barlow www.breeders-registry.gen.ca.us/ light regimes feed for longer periods et al. (1995) examined the growth of species.htm of time than those reared under barramundi larva (Lates calcarifer) Anon. 1998. C-Quest homepage: http:// 12L:12D photoperiod, thereby yield- under different photoperiods, and c-quest.com/fishdata.htm ing higher rates of growth and de- found that individuals 8-20 days Arvedlund, M. and L.E. Nielsen. 1996. velopment. The fact that growth old had significantly higher growth Do the anemonefish Amphiprion under 24L:0D was slower than un- rates with photoperiods of 16-hour ocellaris (Pisces: Pomacentridae) imprint themselves to their host sea der 16L:8D suggests that unless the and 24-hour light. A similar result anemone developing juveniles have a period was also obtained for sole (Soela (: Actinidae)? Ethology of inactivity during darkness, their solea) by Fuchs (1978). 102:197-211. growth is compromised. In contrast, Kiyonon and Hirano Barahona-Fernandes, M.H. 1979. Some Our findings are supported by (1981) reported an optimum growth effects of light intensity and photo- earlier studies of other fish species. rate of black porgy (Mylio macro- period on the seabass larvae A study of the rockfish Sebastes cephalus) with continuous light. (Dicentrarchus labrax (L.)) reared diploproa was reported to have an Tandler and Helps (1985) also re- at the Centre Oceanologique de optimum growth rate with a 16-hour ported this for gilthead sea bream Bretagne. Aquaculture 17:311-321.

22 Naga, The ICLARM Quarterly (Vol. 23, No. 2) April-June 2000 Hoff, F.H. 1993.Marineornamentalfish Hioki, S.,S.KatsumiandY. Tanaka. Gardner, T.R. 1997.Commercialbreeding Fuchs, J.1978.Influencedela Frakes, T. andF.H. Hoff, Jr. 1982.Effect Fautin, D.G.1991.Theanemonefish Naga, TheICLARM Quarterly(Vol. 23,No.2)April-June2000 Duray, M.andH.Kohno.1988.Effects Danilowicz, B.S.andC.L.Brown.1992. Coughlin, D.J.1994.Suctionpreycap- Brons, R.1996.Reproductionandcap- Boehlert, G.W. 1981.Theeffects ofpho- Barlow, C.G.,M.G.Pearce,L.J.Rodgers Society, Oostende(Belgium).Spec. cialization. EuropeanAquaculture culture. Fromdiscoverytocommer- 37(1):34-38. ferrugatus larva intheangelfish, 1990. Developmentofeggsand of thedottybacks.Seascope14:1-2. ture 15:63-74. ( survie delalarveetdujuvenilesole photoperiode surlacroissanceet Aquaculture 29:155-158. anemonefish, survival ofjuvenileandlarval of highnitrate-Nonthegrowthand is not.Symbiosis10:23-46. : whatisknownand Aquaculture 109:311-321. rabbitfish, and survivaloffirst-feedinglarval of continuouslightingongrowth culture 106:141-149. (Gill) and fish species: Rearing methodsfortwodamsel- 1:242-246. ( ture byclownfishlarvae Freshwat. Mar. Aquar. 19(6):48-62. fridmani : tive breedingoftworedsea 79(4):789-794. growth inthefield.Fish.Bull. diploprora ratory growthofjuveniles toperiod andtemperaturesonlabo- (Bloch). Aquaculture138:159-168. venile barramundi feeding periodicityoflarvalandju- toperiod ongrowth,survivaland and P. Clayton.1995.Effects ofpho- Solea solea Amphiprion perideraion and . JapanJ.Ichthyol. D. aruanus and acomparisonwith ) enelevage.Aquacul- Amphiprion ocellaris Siganus guttattus Dascullys albisella P. flavivertex Lates calcarifer Pseudochromis (L.). Aqua- Centropyge ). Copeia Sebastes . . . Job, S.D.andD.R.Bellwood.1996.Vi- Hoff, F.H. 1996.Conditioning,spawn- Tandler, A.andS.Helps.1985.Theef- Schumann, B.1969.Rearingmarine Patzner, R.A.andR.Brandstaetter. 1989. Moe, A.M.Jr. 1992.Themarine Moe, M.1975.RearingAtlanticangel- Miyagawa, K.1989.Experimental McLarney, B.1986.Pioneersinsaltwa- McLarney, B.1985.Pioneersinsaltwa- Marini, F.C. 1996.Mynotesandobser- Lange, J.1989.Thebreedingofdiffer- Kiyonon, M.andR.Hirano.1981.Ef- Juhl, T. 1992.Commercialbreedingof Job, S.,M.ArvedlundandMarnane. Premnas biaculeatus sual acuityandfeedinginlarval Florida, USA.212p. Aquaculture ConsultantsInc., phasis onmarineclownfish. ing andrearingoffish Publ. Eur. Aquacult.Soc.19:238. fects ofphotoperiodandwaterex- 5(5):1-10. fish larvae.SaltWater Aquar. Mag. naco) 5:223-226. 1810. Bull.l’Inst.Oceanogr. (Mo- Rearing of 318 p. Florida, USA. breeder. GreenTurtle Publ aquarium handbook.Beginnerto fish. Mar. Aquarist7(7):17-26. Ethology 80:19-46. anemonefishes andseaanemones. analysis ofthesymbiosisbetween Mag. (January/February):31-33. ter aquariumfish.PartII.Aquacult. Mag. (November/December):38-41. ter aquariumfish.PartI.Aquacult. 4(4):1-4. .J.MaquaCult. vations onraisingandbreedingthe 219-222. Oceanogr. (Monaco)No.5Spec.: Aquarium Berlin.Bull.l’Inst. ent coralfishesintheZoo- Réun. CIESM178:334-336. postlarvae andjuveniles.Rapp.P.-V. rocephalus growth ofblackporgy, fects oflightonthefeedingand anemonefishes. Seascope9:1-4. Austasia Aquacult.11(3):56-59. 1997. Cultureofcoralreeffishes. 48:952-963. Blennius pavo (Basilewsky), . J.Fish.Biol. with em- Mylio mac- ication, Risso, Terver, D.1975.Donnessurl’elevageet Terver, D.1971.Comportementde Young, F.A. 1994.Rearingthegolden Young, F.A. 1990.RearingtheRedSea Young, F.A. 1982.Theyellowhead Wilkerson, J.D.1998.Clownfishes.A Wassink, H.1990.EinegelungeneZucht Walker, S.D.1991.Reproducingthe Young, F.A.state oftropical 1996.The and QLD . Cook University, Townsville 4811 partment ofMarineBiology, James d’ la reproductionnaquarium polymnus poisson-clown ( eloppement embryonnaired’un ponte, reproductionetdev- 48:71-82. rearing systems.Aquaculture hatching tometamorphosisinmass aurata of giltheadseabream( change rateongrowthandsurvival (Pomacentrids). goby, Freshwat. Mar. Aquar. 5:136-137. anemonefish Aquarium 5:50-51. mouthbreeder incaptivity. jawfish. Breedingthemarine p. crocosm Ltd.,Shelburne,VT. 240 ing andnaturalhistory. 1sted.Mi- guide totheircaptivecare,breed- Aquar. Ter. Z.3:126-130. altivelis des MIrakelfisches, Seascope 8:1-3. Atlantic spadefishattheTulsa zoo. Nancy:310-321. Thése d’Etat. en aquarium.UniversitdeNancyI, biologie etauxtechniquesdeslevages 129. Mar. tion. Freshwat. Aquar. 5:126- marine aquariumanimalcultiva- Freshwat. Mar. Aquar. 12:182-184. M. A T. A T. Amphiprion allardi RVEDLUND INSWORTH , Linnaeus;Sparidae)from ). Piscic.Fr. 26:9-23. Elactinus xanthipora (Steindachner, 1903). Amphiprion bicinctus In , M.I.M are from theDe- Contributionala Calloplesiops Amphiprion C Klausewitz C ORMICK Sparus 23 . .

AQUABYTE