Calif. Mar. Res. Comm., CalCOFI Rept., 14 :76-83, 1970.

FEEDING LARVAL MARINE FISHES IN THE LABORATORY: A REVIEW ROBERT C. MAY ’ Scripps Institution of Oceanography La Jolla, Cailfornia INTRODUCTION of this well-studied family of predominantly marine fishes. Interest in rearing marine fishes under laboratory The present review will summarize previous at- conditions has intensified during the past decade, and tempts to feed the larvae of marine fishes in the there is every reason to believe that this trend will laboratory by presenting 1) a list of all food types continue. The larval stages of marine fishes are being used in the studies reviewed, and 2) a general discus- investigated experimentally because of their inherent sion of the major food types. Only reports describing biological interest, their importance in determining the rearing of larvae hatched from eggs in the labora- recruitment in adult fish populations, and their cen- tory are considered here, and reports of rearing in tral place in problems associated with large-scale fish large, uncontrolled bodies of water such as ponds, culture. The greatest difficulty in attempts to rear are excluded. Also excluded, of course, are studies marine fishes through the larval stage has frequently dealing with the rearing of fishes which have no free- been to provide a suitable food-that is, a food which swimming larval stage. A number of papers have ap- the larvae will consume and grow on and which can peared during the 1960’s concerned with the physi- be supplied in sufficiently large quantities. The litera- ology and behavior of marine fish larvae; although ture on rearing marine fishes is scattered, in several they frequently include mention of feeding in the languages, and in some cases not readily accessible, laboratory, most of them have been omitted here since and a comprehensive review emphasizing feeding has the methods used are usually described in greater not appeared until now. This review summarizes the detail in other papers more specifically concerned attempts to feed larval marine fishes in the laboratory with rearing techniques. which have been described in the literature from 1878 One of the drawbacks of reviews of this kind is that through 1969, in the hope that such a review will aid they necessarily oversimplify the original work and persons who undertake similar work in the future. thus may be misleading if not used properly. When Several previous reviews contain information rele- dealing with reports-many of which are sketchy al- vant to the problem of feeding larval fishea. Hert- most to the point of uselessness-of experiments con- ling’s treatise on rearing marine fishes (Hertling, ducted under vastly different conditions with a 1932) includes a section on feeding, but investigators variety of species, comparisions are difficult and now will find his discussion of little help, partly be- generalizations risky. If a particular food type was cause at the time he was writing, relatively little work used in the past without success, it is still possible had been done on rearing beyond the yolk-sac stage that it will prove useful for another species of fish or under laboratory conditions. Morris (1955) confined for the same species under different conditions. Also, his discussion of nutrition in marine fish larvae to a several different approaches to the problem of feed- consideration of Putter’s theory that dissolved or- ing may prove equally fruitful. A case in point is the ganic matter is an important source of nutrition for rearing of the sole, Solea solea: Fabre-Domergue and fishes, an hypothesis which has so far not been de- Bihtrix (1905) reared sole using a flagellate as first finitively tested in larval fishes. Morris (1956) de- food and, later, wild plankton from which the larvae scribed rearing experiments with 17 species of selectively preyed upon other fish larvae and ignored California fishes and included a lengthy discussion of copepod nauplii ; nevertheless, Fluchter (1965) ob- various food types. Hirano and Oshima (1963) re- tained excellent results by feeding larvae of this viewed methods of feeding the larvae of a variety of species only artemia nauplii. marine animals, including selected examples of at- The emphasis of the present review on feeding is tempts to feed larval fishes, and supplied valuable not meant to imply that other factors are not im- data on the sizes of various potential food organisms. portant for the successful rearing of larval fishes, or Shelbourne (1964) described his extensive work with that larval survival in the studies reviewed was solely the plaice and gave an account of the historical de- dependent upon the choice of food. Providing a suit- velopment of marine fish culture. Blaxter (1965) able food is a sine qua non in any rearing attempt, discussed the feeding ecology of herring larvae, inte- but a host of other significant factors are operating grating data from the laboratory and the field, and in the complex artificial environment of the larvae. Blaxter and Holliday (1963), reviewing the biology The importance of such factors is evident, for ex- of the clupeids, summarized information on the larvae ample, in Blaxter’s (1968a) observation that although 1 National Marine Fisheries Service Fellow. This work was done over 90% of the herring larvae in his experiments at the National Marine Fisheries Service Fishery-Ocean- ography Center, La Jolla, California. were feeding well, only 25-35% survived yolk-ab- REPORTS VOLUME XIV, 1 JULY 196s TO 30 JUNE 1969 77 sorption. Also, it is not uncommon to find dead larvae TABLE 1-Continued with food in their guts. A discussion of factors other Foods Used in Attempts to Rear the larvae of Marine Fishes than food is beyond the scope of the present review, and persons interested in the details of the studies Food References reviewed here must consult the original papers. Bacillariophyceae Coscinodiscus coneinnus------Hertling, 1932 LIST OF FOOD TYPES Coscinodiscus radiatus -... - - - - Gross, 1937 Skeletonema costatum- - - - -.- - - Blaxter, 1962; Forrester, 1964; Gross, Table 1 lists the food types which have been used 1937; Kurata, 1956, 1959 in all papers included in the review. For con- Thalassiosira sp _.__ ___.__ Gross, 1937 Lauderia borealis- - __ -.- __ __.- Blaxter, 1969 venience the living organisms other than wild plank- Rhizosolenia sp - ._._.___David, 1939 ton have been arranged taxonomically under four Chaetoceros sp David, 1939 main headings : Protista: planktonic forms; Protista: Biddulphia mobiliensis------Hertling, 1932 Ditylum briahtwellii- - - - - __ -.. Blaxter, 1969 nonplanktonic forms; Metazoa: planktonic forms; Nitzschia closteium .__._.- ._._ Drcutt, 1950 and Metazoa: nonplanktonic fornas. In a few cases Nitzschia sp Bishai, 1961; Blaxter. 1962: Budd, 1940 “Diatoms”------.- - __ ------David, 1939; Rubinoff, 1958 the designations planktonic and nonplanktonic are somewhat arbitrary. Nonliving foods and “prepared Eumycophyta Ascomycetes aquaria” have been placed under the heading Miscel- “Yeast”...... ~. ._._ .._ .____ Klirna et al.. 1962; Morris, 1956 laneozts. Taxonomy uncertain “Natural and cultured phyto- plankton” - - -..------Klima et al., 1962 TABLE 1 “Raw cultures of phytoplankton” Dannevig, 1948 “Cultures of single species of green Foods Used in Attempts to Rear the larvae of Marine Fishes algae and naked flagellates”--- Dannevig. 1948 - “Phytoplankton”_ -... . - - -.- - - - Bishai. 1961; Ivanchenko & Ivan- chenko, 1969 Food References “Plankton algae”. - - -.-. _. __ -. - Runnstrom. 1941 --- “Algae”. .. -. .. -. .. . ------Fabre-Domergue & Bibtrix, 1897 “Monadinen”. .. - -... -. .-. - - -. Kotthaus, 1939 WILD PLANKTON “Vegetarian diet” .___.-.-. -. _. - Soleim, 1942 Antony, 1910; Blaxter, 1968, 1969; Blaxter and Hempel, 1961; Budd, 1940; PROTISTA: Cunningham, 1893-95a & b; A. Dan- NONPLANKTONIC FORMS nevig, 1948; H. Dannevig, 1897; Dan- Cyanophyta nevig & Hansen. 1952; David, 1939; “une nostocacee” - -.- __ ._. .. -. . Fabre-Domergue & BiBtrix, 1897 Fabre-Domergue & Bibtrix, 1897,1905; Garstang, 1900; Ivanchenko & Ivan- Chlorophyta chenko, 1969; Kramer & Zweifel, 1970; Enleromorpha sp.. ___ ..__._.. Kotthaus, 1939 Lebour, 1925; Meyer, 1878; Mito et al., 1969; Morris, 1956; Okamoto. 1969; Chrysophyta Orcutt, 1950; Richards & Palko, 1969; “Filamentous brown diatom”- __ Fabre-Domergue & Bidtrix, 1897 Schach, 1939; Schumann, in Bardach, 1968; Yamamoto & Nishioka, 1952 Ciliophora Ciliata PROTISTA: PLANKTONIC Euplotes sp Fabre-Domergue & Bidtrix, 1897 FORMS Philaster digitiformis. ------Fabre-Domergue & Bidtrix, 1897 Chlorophyta Stylonychia sp Kasahara et al.. 1960 Loxophyceae Platymonas subcordiformis-----. Orcutt, 1950 Taxonomy uncertain Halosphaera minor __.______.Blaxter, 1969 “Filamentous algae”- - -..__. _.- Delmonte et al., 1968 Chlorophyceae “Cultured infusorians”- - -.- - - - - Fabre-Domergue & BiBtrix. 1897 Chlamydomonas sp _._... Bishai, 1961; Blaxter, 1962,1969; Gross, 1937: Qasim, 1955, 1959 METAZOA: Dunaliella primoleeta- -.-. -. -. . Blaxter, 1969 PLANKTONIC FORMS Dunaliella salina-- __ - __ ------. Fabre-Domergue & BiBtrix, 1905 Aschelminthes Dunaliella sp Blaxter, 1962; Budd, 1940; Delmonte Rotifera et al., 1968; Morris, 1956 Brachinous plicatilis--- __ _.-.- Hirano, 1969; Mito et al., 1969; Oka- Chlorella stigmata.. - - - - ______. Qasim, 1955 moto. 1969 Chlorella sp Blaxter, 1962 Mollusca Stichococcus sp Morris, 1956 Gastropoda Crepidula sp. “larvae” ____ - - -. Rubinoff, 1958 Pyrrophyta Bulla aouldiana ueligers___._. . Lasker et al., 1970 Cryptophyceae Haminoea vesicula veligers- - -. Lasker et al., 1970 Cryptomonas maculata------. Blaxter, 1969 Nananaz inermis veligers-----. Lasker et al., 1970 Desmophyceae “Nacktschnecken” (7 nudi. Prorocentrum micans- - - - -.- - -. Blaxter, 1969; Gross, 1937, Lasker et al., branch) larvae.____._._____ Kotthaus, 1939 1970; Qasim, 1955 Pelecypoda Mytilus californianus “larvae” Morris, 1956 Dinophyceae Mytilus sp. trochophores-- - __ . Blaxter & Hempel, 1961; Okamoto, Gymnodinium splendas------. Lasker et al.. 1970 1969; Schach, 1939 Oxyrrhis marina.. Morris, 1956 Mytilus sp. “larvae”. __.___.. Dannevig, 1948; Kotthaus, 1939; Kur- Ozyrrhis sp Kasahara et al., 1960 ata, 1956 Protoceratium reticdatum- - - - - . Lasker et al, 1970 Ostrea edulis “larvae”-. _____.Dannevig, 1948 Fragilidium heterolobum____ - - - . Lasker et al.. 1970 Crassostrea gigas “larvae”.--. . Hirano, 1969 “Oyster” trochophores ------. Okamoto, 1969 Crysophyta Xanthophyceae Annelida Olisthodiscus sp Blaxter, 1969 Polychaeta Chrysophyceae Chone teres trochophores-----. Kurata, 1959 Isochrysis galbana ______-. Qasim, 1955, 1959 Pomatoceros sp. “larvae” ____ -. Dannevig, 1948 Monochrysis lutheri ______-. Forrester, 1964 Nereis sp. eggs ______._____Cunningham, 1893-95b Chromulina pusilla------. Qasirn, 1955 “Minced worms” ______.Cunningham. 1893-95a 78 CALIFORNIA COOPERATIVE OCEANIC FISHERIES IKVESTIGATIONS

TABLE 1-Continued TABLE 1-Continued Foods Used in Attempts to Rear the Larvae of Marine Fishes Foods Used in Attempts to Rear the Larvae of Marine Fishes ~- Food References Food References

~~

METAZOA: Echinodermata PLANKTONIC FORMS Echinoidea -Continued Dendraster excentricus eggs or Arthropoda larvae--.-----.-.-.-..---- Morris, 1956 Crustacea Strongylocentrotuspurpuratus Arlemia salina nauplii- - ._._._ Bishai. 1961; Blaxter, 1962,1968; Blax- eggs or larvae ___....______Budd, 1940: Morris, 1956: Orcutt, 1950 ter & Hempel, 1961; Budd, 1940: Tripneustes esculentus eggs or Chirinos de Vildoso & Chuman, 1964; larvae--.-.-.-..------.- Richards & Palko, 1969 Dannevig, 1948; Dannevig & Hansen, Arbacia sp. eggs or larvae--.-- Deuel et al., 1966; Rubinoff. 1958 1953; Delmonte et al., 1968; Fahey. “Fertilized sea urchin eggs”--- Fujita, 1965 1964; Fishelson, 1963; Fliichter, 1965; Asteroidea Forrester, 1964; Fujita, 1957, 1965, Asterias sp. eggs .._._._._.... Blaxter, 1962 1966: Gross, 1937: Hirano, 1969; Joseph Urochordata & Saksena. 1966; Kasabara et al., 1960; Ascidian larvae._.._._._.._._ Fabre-Domergue & BiBtrix, 1897 Klima et al., 1962; Kramer & Zneifel, Chordata 1970; Kurata. 1956, 1959; McHugh & Fish larvae ._...._.___.._._._Fahre-Domergue & BMtrix, 1905 Walker, 1948; McMynn & Hoar, 1953: Mito et al., 1969; Molander & Molander- METAZOA: Saedmark, 1957; Morris, 1956; Oka- NONPLANKTONIC FORMS moto, 1969; Orcutt, 1950; Qasim, 1955. Aschelminthes 1959; Richards & Palko, 1969: Rollef- Nematoda sen, 1939; Rubinoff, 1958; Schumann, Anguillicula sp. ._..._..._..._ Blaxter. 1962 in Bardach, 1968: Shelhourne. 1964: Shojima. 1957; Soleim, 1942 RIISCELLANEOGS “Cladocera” ------.. . -. -. - - - Gross, 1937 Prepared aquaria...... ~.__ ._._._Bishai, 1961; Delmonte et al., 1968; Daphnia pulex eggs .__...____Mchlynn & Hoar, 1953 Heuts, 1947; Klima et al., 1962: Mc- Daphnia sp Blaxter, 1965; Biickmann et al.. 1953 Hugh & Walker, 1948 “Copepods”. .-. -. -...... Bishai. 1961; Biickniann et al.. 1953; Finely ground trout food .__...__._Richards & Palko. 1969 Fishelson, 1963; Gross, 1937; Hirano, Powdered fish foods_.....__._._._ Rubinoff, 1958 1969 Commercial fish-fry foods...... Blaxter, 1962: Delmonte et al.. 1968; “Copepod nauplii“.. .-...... Kasahara et al., 1960; Kotthaus, 1939 Klima et al., 1962 Tigriopus californicus nauplii-. Fahey, 1964 Cooked chicken egg yolk __.___._._Biickmann et al., 1953: Fabre Domergue Tigriopus fuluus nauplii _._..- - Bishai, 1961; Budd, 1940; Orcutt, 1950 & BiBtrix, 1897; Fishelson. 1963; Fujita, Tigriopus fulvus adults. ___... Blaxter, 1965, 1968; Morris, 1956 1966; Heuts, 1947; Ivanchenko &Ivan- Tigriopus sp. “young stages”- Blaxter, 1962 chenko, 1969; Klima et al., 1962; Kur- Tisbe sp Blaxter, 1962 ata, 1959; Morris, 1956; Nikitinskaya, Balanus balanoides nauplii.. .. Blaxter, 1962, 1968; Dannevig, 1948; 1958 Qasini, 1955. 1959; Shelbourne, 1964; Water-soluble vitamin compounds- Klima et al., 1962 Soleim. 1942 Liver of shore crab, Carcinus maenas Bishai, 1961

Balanus glandula nauplii L... Morris, 1956 Minced shrimp and crab meat..-.- Kurata, 1959 Balanus amphitrite albicostatu. Crushed mussel.-...... Fabre-Domergue & BiBtrix. 1897 nauplii.. .-...... -. .. Hirano, 1969; Kasahara et al., 1960 Homogenates of Mytilus, periwinkle, Neomysis japonicus “young”. . Kasahara et al.. 1960 Fucus and kelp ...... Iranchenko & Ivanchenko, 1969 Liver-skim milk __...._..._._..._McMynn & Hoar, 1953 Human blood ...... _._._...._...Klima et al., 1962

DISCUSSION OF THE MAJOR FOOD TYPES ing experiments, often list the dominant organisms in the plankton. Some also give information on which Wild plankton. One of the most widely used sources plankters were eaten by the larvae, based either on of food for larval fishes has been plankton taken from analyses of gut contents or on observations of feed- natural waters, referred to here as wild plankton. ing behavior. Copepods, copepodites and copepod Wild plankton is usually collected by tows with a nauplii frequently dominate in wild plankton and plankton net, although Schumann (work described in often constitute the main food of the larval fishes Bardach, 1968) collected plankton at night with a (Garstang, 1900 ; Lebour, 1925 ; Schach, 1939 ; Blax- submerged pump to which plankters were attracted by ter, 1969), but many other types of organisms may a light, and concentrated the plankton with a series of be present (cf. Blaxter and Hempel, 1961 ; Blaxter, filters. Prior to feeding the plankton to the larvae, it 1968a), and herring and plaice larvae are reported to is common practice to strain it through netting of a have selectively fed upon larval molluscs in wild mesh size which passes only those organisms which the plankton (Meyer, 1878 ; Dannevig, 1897). Blaxter larvae can feed upon. This performs the additional (1969) had more success with offshore than with in- function of removing most potential predators from shore plankton in rearing experiments with larval the plankton. Meyer (1878), however, noted poor sardines. survival and growth among herring larvae which were given plankton strained through cloth and found that Wild plankton has proved to be an excellent source normal growth was resumed when unstrained plank- of food for larval fishes in the laboratory, as might ton containing large numbers of copepods was ad- be expected since it contains the same food which fish ministered. It would seem obvious that as the lame larvae feed on under natural conditions. Plankton grow the size range of the plankton used as food collected in the sea was the mainstay of the successful should be increased. The plankton supplied to larval early attempts to rear marine fishes (Meyer, 1878; fishes usually contains a mixture of organisms, and Dannevig, 1897 ; Garstang, 1900 ; Fabre-Domergue authors who describe the use of wild plankton in rear- and Biktrix, 1905; Anthony, 1910) and is still being REPORTS VOLUME XIV, 1 JULY 196s TO 30 JUNE 1969 79 used in rearing for experimental purposes (Schu- could be reared from hatching through metamorphosis mann, in Bardach, 1968 ; Blaxter, 1968, 1969 ; Ivan- with nauplii of the brine shrimp, Artcmia salina L., chenko and Ivanchenko, 1969 ; Richards and Palko, as the only food. This is an extremely convenient 1969 ; Kramer and Zweifel, 1970). source of food, since the nauplii hatch readily when However, despite the proven effectiveness of wild the dried eggs are placed in water, and the eggs can plankton as a food source, its use has serious disad- be stored for long periods and used when needed. Un- vantages which make it unsuitable in many instances. fortunately, not all fish larvae are able to take Artcnzia Among these disadvantages are : 1) the seasonality nauplii as a first food because of their relatively large and highly variable quality and quantity of wild size (length 0.4 mm, width with appendages folded plankton ; 2) the dependence of plankton collection on 0.35 mm; from Blaxter, 1962). Hirano and Oshima weather conditions ; 3) the time-consuming nature of (1963) pointed out that larvae of different fish species collecting plankton ; 4) the mixture of food types in begin taking Artcnaia nauplii at different sizes. For the plankton which makes controlled feeding experi- example, the puffer, Fiigzi rzcbripcs, eats Artenaia ments difficult; and 5) the difficulty of removing from nauplii when it is 4.1 mm long ; Mylio macrocephalus the plankton all organisms which may prove harmful and the flatfish, Liopsctta oOsczira at 5 mm ; the half- to the fish larvae. Because of such drawbacks, much beak, Henairamphzis sajori, at 6-6.5 mm; and the Paci- effort has been expended in the search for larval-fish fic herring, Clupca pallasii, at 9-10.5 mm (Hirano foods that can be cultured in the laboratory. The long and Oshima, 1963). Fishelson (1963) reported that list of food types presented in Table 1 testifies to Blcnniiis pavo of 3.5 mm total length will Consume this fact. Artenzia nauplii, and Fliichter (1965) found that Phytoplankton. Of the many attempts described in Xolca solea larvae 3 to 4 days old, Le., ea. 5 mm total the literature to feed phytoplankton to larval fishes length, began to eat this food. These differences are (Table l), very few have met with any success. In undoubtedly related to differences between species in several experiments various types of phytoplankters morphometry and mouth size (see Schumann, in Bnr- were ingested by fish larvae, yet the larvae survived dach, 1968), and perhaps also to the use of different no longer than starved controls (Gross, 1937 ; Orcutt, strains of Artcmia with different sized nauplii. Roseii- 1950 ; Qasim, 1955, 1959 ; Bishai, 1961 ; Blaxter, thal and Hempel (1970) wrote that among young her- 1969) ; this suggests that at least certain types of ring, the success of prey-catching maneuvers WHS phytoplankters are nutritionally inadequate for larval higher with Artcmia nauplii than with wild plankton, fishes. Xchach (1939) found that yolk-sac herring lar- and they attributed this to the greater visibility of vae kept in water thick with phytoplankton acquired Artenzia (Blaxtcr, 196813) and to its sluggish escape a “green mass” in their guts, presumably from ingest- response. ing phytoplankton, but these larvae did no better than Certain flatfish larvae appear to do very well on an larvae kept in clear water, and both groups began exclusive diet of Artenzia nauplii. Xhelbournc (1964) feeding on zooplankton 4 days after yolk-absorption. reported good survival (66‘;: ) through metamorphosis for plaice larvae feeding on Artcmia alone, and In the first successful attempt to use phytoplankton Fliichter (1965) fed sole larva(. on Artcmia and ob- as a larval fish food, Fabre-Domergue and Bi6trix tained 8031 survival through metamorphosis, and (1905) found that newly-hatched sole larvae, Solea growth comparable to that of larvae in the sea. solea, ate Dzinaliclla salina cultured from salt marshes Artcmia nauplii alone have also been used success- but soon switched to carnivorous feeding. IIertling fully to feed larval Pzindicliis hctcr-oclitus (Joseph (1932) stated that the armed bullhead, Agonzis cata- and Saksena, 1966), Fiindzclzts nzajalis (Fahey, 1964), phractiis, could be reared by feeding the early larvae Azilorlzynchzis flavidiis (Morris, 1956), Sebastes pac- on diatoms, and Runnstrom (1941) claimed to have hyccphaliis nigricans (Fujita, 1957), two cottid kept herring larvae alive for 6 weeks on “plankton species (Morris, 1956), four species of artherinids algae,” but these authors supplied few details. Kasa- (McHugh and Walker, 1948 ; Morris, 1956 ; Chirinos hara et al. (1960) fed early larvae of the black porgy, de Vildoso and Chuman 1964), and Fzigii parclalis Mylio macrocephalzis, on the naked dinoflagellate (Shojima, 1957). They have frequently been used to Oxyrrhis but soon changed over to animal foods. supplement other food types (e.g. Dannevig iind Hiin- Lasker et al. (1970) gave the only detailed description sen, 1952 ; Blaxter and Hempel, 1961 ; Blaxter, 1962, of rearing for prolonged periods using phytoplankton 1968a; Fishelson, 1963) or to feed older larvae (e.g. alone. They reared the northern anchovy, Engraidis Danncvig, 1948 ; Kasahara et al., 1960; Schumann, in mordax, for 19 days (when the experiments were pur- Bardach, 1968 ; Okanioto, 1969 ; Kramcr and Zweifel, posely terminated) with good growth and survival, 1970). using the naked dinoflagellate Gymnodinium splen- Riley (1966) has shown in a controlled experiment dew as the sole food, and in one experiment kept a that the survival and growth of plaice larvae feeding larva alive for 34 days on this phytoplankter. Growth on Artcnaia nauplii are markedly affected by the and survival were somewhat better, however, when amount of food available. He found that surplus food molluscan larvae were supplied along with the increased the survival beyond first feeding ; but very Gymnodinizcm. high feeding levels were detrimental to the larvae Artcmia nauplii. Perhaps the greatest breakthrough after feeding had become established, since they in feeding techniques was the discovery by Rollefsen caused fouling of the water and promoted the growth (1939, 1940) that the plaice, Pleuronectes platessa, of bacteria and ciliates. Fliichter (1965) recom- 80 CALIFORNIA COOPERATIVE OCEANIC FISHERIES ISVESTIGATIONS mended extremely high concentrations of nauplii (20,- usefulness as larval fish food (Blaxter, 1962, 1968a). 000 per liter) for first-feeding sole larvae. The neces- Blaxter (1965, 1968b) found that herring larvae re- sity of high concentrations of food for young larvae quired more light to feed on Balanus nauplii than on is also strongly suggested by Rosenthal’s (1969b) Arteniia nauplii, presumably because the former are finding that first-feeding herring larvae were success- smaller and more transparent. ful in only 1%to 10% of their strikes at prey. Since nauplii of calanoid copepods are frequently A rather substantial body of evidence indicates that the dominant organisms in the guts of larval fishes Artenaia nauplii alone will not sustain the growth and caught at sea (e.g., IJebour, 1920), they would appear survival of clupeid larvae over long periods. Dannevig to be a natural choice as a food for use in laboratory and Hansen (1952) found that older larvae of the fish-rearing. But copepod nauplii have never been CUI- Atlantic herring, Clupea harengus, did not grow on an tured on a large scale for the purpose of feeding larval exclusive diet of Artemia. Blaxter and Hempel (1961) fishes, probably because of the technical difficulties of and Blaxter (1962) also had little success using only producing sufficient numbers. Kotthaus (1939) har- Artemia to feed C. harengus, and Blaxter (1968a) vested nauplii which hatched from eggs carried by was unable to rear herring beyond 25 mm on Artemia. adult copepods collected in plankton tows, but he McMynn and Hoar (1953) and Kurata (1959) reared worked on a rather small scale. Kasahara et al. (1960) the Pacific herring Clupea pallasii only to 10.2 and listed copepod nauplii as one of the foods they used 14.5 mm, respectively, on Artcmia alone. An explana- but did not say how the nauplii were obtained. tion for this phenomenon may be that the digestive Morris (1956) stated that nauplii of the spray-pool system of the larval clupeid is incapable of fully di- harpacticoid Tigriopus sp. are thigmotactic and hence gesting the nauplii of Artemia. Rosenthal’s (1969a) not available to larval fishes, and most recorded at- observation that Artemia nauplii are much more re- tempts to use them as food have met with no success sistant to digestion by larval C. harengus than are (Budd, 1940 ; Orcutt, 1950 ; Bishai, 1961 ; Blaxter, copepods supports this explanation. Attempts to feed 1962). Nevertheless, Fahey (1964) mentioned, without Artemia nauplii on phytoplankton in order to make further elaboration, that he reared Bairdiella chry- them more nutritious (Dannevig and Hansen, 1952 ; sura, Alosa pseudoharengm, and Mugil caphalus using Morris, 1956; Blaxter, 1962) appear to have met with mass-cultured first and second naupliar stages of T. little success, although Rosenthal (1969b) fed herring californicus as food. Certain other crustaceans such as larvae of 15 mni total length and larger on Artemia young mysids (Kasahara et al., 1960) and adult Ti- which had been reared on cultured Dunaliella to ea. griopits (Morris, 1956 ; Blaxter, 1965, 196th) have 1.8 mm in length. Despite the evidence for the nutri- been used to feed older larvae. tional inadequacy of Artemia nauplii for clupeid lar- Moll~sclarvae. The larvae of certain molluscs are vae, Blaxter (1962) found that 10-12 mm herring attractive food sources because of their availability which had previously fed on Artenaia nauplii survived and small size (most under 0.2 mm, many under 0.1 up to twice as long without food as larvae which had mm-see Loosanoff and Davis, 1963; Hirano and fed on Balaniis nauplii. Oshima, 1963) and have been used successfully several Certain technical problems are associated with the times, especially to feed very young larvae. Most such use of Artenzia nauplii. Care must be taken to avoid experiments have employed the larvae of bivalve mol- placing eggshells in rearing containers along with the luscs, which generally hatch as trochophores and later nauplii, as they may be eaten by fish larvae in place develop into veligers. Kurata (1956) found mussel of nourishing food and the larvae may starve as a larvae an excellent food for early larvae of the flatfish result (Morris, 1956). Rosenthal (1969a) observed Liopsetta obscura, and Okamoto (1969) gave mussel one herring larva which took almost nothing but Ar- and oyster trochophores to early red sea bream, Pag- temia eggs. Blockage of the gut by Srtemia eggshells ribs major, larvae. Morris (1956) describes a procedure has been reported in young seahorses (Herald and for obtaining eggs from the California mussel, Mytilzcs Rakowicz, 1951). As Artemia nauplii grow, they may californianus. Dannevig ( 1948) used oyster larvae suc- become too large for the fish larvae to eat; they also cessfully to feed smaller larvae of several fish species, become less nutritious as their yolk is consumed, a and Hirano (1969) reared Mylio niacrocephalz~susing change which is indicated by transition from a deep oyster larvae as the first food. Gastropod larvae, which orange to a whitish coloration (Morris, 1956). Morris in most cases hatch as veligers, have been used much stated that fish larvae which fed on older Artenzia less frequently than bivalve larvae. Lasker et al. nauplii “did not prosper,” and he recommended that (1970) found that the veligers of three species of opis- uneaten nauplii be removed from rearing tanks each thobranch gastropods were too large to be taken by day. first-feeding anchovy larvae but that they increased the growth rate and survival of older larvae which Other crustaceans. Nauplii of the barnacle Ealanzcs had fed first on a dinoflagellate. May (unpublished sp. are smaller than Artemia nauplii (Hirano and data) reared the atherinid Leuresthes tenuis through Oshima, 1963) and have been used to feed larval C. metamorphosis using veligers of the gastropod Bulla harengzis (Soleim, 1942 ; Dannevig, 1948 ; Blaxter, 1962, 1968a), Pleuroncctes platessa (Shelbourne, gouldiana as the sole food, with a survival of 16.7 percent through metamorphosis. 1964), Elennitis pholis and Centronotus gunnellus (Qasim, 1955, 1959) and Mylio macrocephalus (Kasa- Other living foods. Kasahara et al. (1960) fed the hara et al., 1960; Hirano, 1969). Barnacles have a hypotrichous ciliate Stylonichia sp. to larval Mylio short spawning season, however, and this limits their niacrocephalus 3-3.5 mni in length. (It is interesting REPORTS vor,ui\m SIV, 1 JULY I~GSTO 30 JUNE IOGO 81 in this connection that Kotthaus (1939), Schach Pngrzcs major the following succession of foods : oyster (1939), and Bishai (1961) considered hypotrichous and mussel trochophores, Brachionzcs plicatilis, bar- ciliates as undesirable contaminants in rearing ex- nacle nauplii, Artemia nauplii, zooplankton, annelid periments with herring and even suggested that they larvae, shrimp meat, and finally fish meat-the ad- may have attacked and killed larvae.) The use of the ministration of each new food type overlapping for rotifer Brachion~tsplicatilis to rear larval fishes has several days that of the previous type. been reportrd recently by Japanese workers (Hirano, Prepared aquaria. In some rearing attempts work- 1969 ; Mito ct al., 1969 ; Okamoto, 1969) who offered it ers have allowed a flora and fauna to develop in the to first-feeding or very young larvae. rearing containers prior to introduction of the fish Kurata (1959) found that first-feeding Clicpca eggs or larvae. These “prepared aquaria” usually pallasii larvae took trochophores of the sabellid poly- contain attached algae, phytoplankton, Zooplankton, chaete Chonc teres in preference to Arfenzia nauplii, and ciliates. Growth of the biota has been encouraged which were larger ; but he considered C. teres as an un- by adding fertilizers and vitamins (Delmonte et al., suitable food source, because its spawning season was 1968) and by placing the tanks in direct sunlight very qhort and the trochophores settled quickly. Cun- (Schach, 1939 ; Heuts, 1947 ; Bishai, 1961). Unfor- ningham (1893-95b) fed Nereis eggs to larval plaice tunately, authors who describe such aquaria rarely with very limited success. specify exactly what organisms are in them, and it Severai workers have tried echinoderm larvae as is impossible to tell from their accounts which items larval fish foods. Morris (1936) reared the sciaenid served as food for the fish larvae. After the fish larvae Geneonynzzts lineatiis beyond yolk-absorption by feed- hatch, specific food items are usually added to pre- ing first with eggs and larvae of the sand-dollar Den- pared aquaria, such as egg yolk (Heuts, 1947) or drasfer and later switching to Artemia nauplii. In Arfenzia nauplii (McHugh and Walker, 1948). The their successful rearing trials Rubinoff (1958), Fujita flora in prepared aquaria seems to maintain chemical (1965) and Richards and Palko (1969) offered sea equilibrium in the water by stabilizing pH and 02 urchin ~ggsor larvae (along with other foods) to tension and taking up fish metabolites. This role of larval fishes but did not state whether this food was green plants was appreciated by some of the early taken to a significant extent. investigators : Garstang (1900) placed one of his Brixun (1949) cited the common use of nematodes rearing containers in direct sunlight and added green for feeding freshwater tropical aquarium fishes and algae (Ectocarpzcs) “as a means of aerating and puri- suggested that they might be a suitable food for marine fying the water,” and Fabre-Domergue and Biktrix fish larvae. But Blaxter (1962) tried the nematode (1902) used algae to assure “d’zine part l’oxyge‘na- Angiiilliczila sp. as a food for herring larvae without tion continue du nzileu, d’azifre part l’e‘limination success, and Korovina (1962), working in freshwater, d’zine proportion de de‘chcfs nzcisibles impossible Ci reported that the hard skin and sharp extremeties of e’vifer.” Shelbourne (1964) incorporated Enteronaor- larval nematodes (Panagrelics redivivzis) injured the plza into closed systems of sea-water circulation for digestive tract of Coregonits larvae and killed them. the specific purpose of stabilizing water chemistry. Riley (1966) found free-living nematodes in the de- Schach (1939) and Bishai (1961) siphoned food tritus on the bottom of some of his tanks which were organisms into their rearing containers from tanks occasionally taken by plaice post-larrae. which were essentially prepared aquaria, containing Algae growing on the walls of rearing containers phytoplankton and copepods and placed in sunlight. was eaten by advanced herring larvae (Kotthaus, Kotthaus (1939) and Schach (1939) both had En- 1939) and by 8-day-old gobies (Delmonte et al., 1968), feromorpha growing in their rearing containers, and and “slime” on tank walls was browsed off by plaice Kotthaus achieved success only in containers which larvae in sufficient quantity to color the gut brown in received some direct sunlight during the day. larvae which had a restricted supply of Artenzia nau- Nonliving foods. Food types in this category are plii (Riley, 1966). attractive because of their great convenience, but they Fabre-Domergue and BiBtrix (1905) found that present the difficulty that uneaten food accumulates sole larvae soon began to feed on other larval fishes, on the bottom of the rearing container and decays including especially sprat larvae but also larvae of yapidly, fouling the water. Anthony (1910) con- their own species. Schumann (in Bardach, 1968) sidered artificial foods unsuitable due to the decay attributed a 50% reduction in the number of Pacific problem, and Fabre-Domergue and Biktrix (1905) mackerel larvae, Scomber japonicus, during the first believed that running water was necessary when arti- 9 days after hatching, to intraspecific predation ; he ficial foods were used. By far the most widely used also found that Pacific barracuda fry, Sphyraena ar- non-living food has been cooked chicken-egg yolk gentea, are exclusively piscivorous at a length of made into fine particles. Fabre-Domergue and Biktrix 4.5 mm. (1897) were apparently the first to try this food source, and Heuts (1947), Fishelson (1963), Fujita Series of food types. Some Japanese investigators (1966), and Ivanchenko and Ivanchenko (1969) have have developed rather complicated feeding schedulcs used it successfully. Egg yolk has served primarily as in which progressively larger food types are offered to the growing fish larvae, there being periods of a food for newly-hatched larvae, and larvae are soon overlap to eliminate abrupt transitions from one food given living food such as wild plankton (Ivanchenko type to the next (Kasahara et al., 1960; Hirano, 1969; and Ivanchenko, 1969) or Arfemia nauplii (Fishelson, Okamoto, 1969). For example, Okamoto (1969) fed 1963 ; Fujita, 1966). Nikitinskaya (1958) , Kurata 82 CALIFORNIA COOPERATIVE OCEANIC FISHERIES INVESTIGATIONS (1959), and Ivanchenko and Ivanchenko (1969) were Budd, P. L. 19-10. Development of the eggs and larvae of six California fishes. Calif. Dept. Fish Game, Fish Bull., (56) : able to keep pallasii alive only for short pe- Cluepea 50 PP. riods on egg yolk alone. Heuts (1947) fed egg yolk to Chirinos de Vildoso, A,, and E. Chuman. 1964. Notos sobre stickleback larvae in prepared aquaria, but it is not el desarollo de hueros y larvas del pejerry Odontesthes clear how long he continued to administer the yolk. (Austromenidia) regia regia (Hiimboldt). Bol. Inst. Mar. Peru, 1: 331. (English summary.) Buckmann et al. (1953) and Fishelson (1963) observed Cunningham, J. T. 1893-95a. The life-history of the pilchard. that particles of egg yolk were consumed by larvae J. Mar. Biol. Ass. U.K., 3: 148-153. only when the particles were moving in water cur- -. 1893-95b. Experiments on the rearing of fish larvae in rents, such as are set up in aquaria by a stream of air the season of 1894. J. Afar. Biol. Ass. U. K., 3: 206-207. Dannevig. A. 1948. Rearing experiments at the Flfidevigen bubbles. Korovina (1962) noted that egg yolk, espe- seafish hatchery 1943-1947. J. Cons. Int. Ezplor. Mer, 15: cially in excess, tended to pollute the water during 277-283. rearing trials in fresh-water with Coregonus larvae. Dannevig, A., and S. Hansen. 1952. Faktorer av betydning for Delmonte al. (1968) successfully fed newly- fiskecgenes og fiskeyngelens opprekst. Fiskeridir. Skr., Ser. et HavundersBk. 10( 1) : 36 pp. (English summary ; English hatched goby larvae in prepared aquaria on com- translation of pp. 6-16, the section on herring : translation mercial fish-fry foods, although they noted a tendency no. 415, Marine Laboratory, Aberdeen, Scotland.) for the uneaten food to decay; Blaxter (1962) and Dannevig, H. 1897. On the rearing of the larval and post-larval Klima et al. (1962) had no success with such foods. stages of the plaice and other flat fishes. Rep. Fish. Bd. &'cot., 1896, Pt. 3 : 175-193, pl. IV. Bits of meat (squid flesh, crab meat, chopped fish, David, L. R. 1939. Embryonic and early larval stages of the etc.) are unsuitable for young larvae (Kurata, 1959; grnnion, Leuresthes tenuis, and of the sculpin, Scorpaena Bishai, 1961), but very advanced larvae commonly guttata. Copeia, (2) : 75-81. have eaten such items (eg. Dannevig, 1948; Kasahara Delmonte, P. J., I. Riibinoff and R. W. Rubinoff. 1968. Labor- atory rearing throiigh metamerphosis of some Panamanian et al., 1960; Blaxter, 1965, 1968; Hirano, 1969). gobies. Copeia, (2) : 411412. Blaxter (1965), however, found that the feeding rate Denel, D. G., J. R. Clark and A. .J. Mansueti. 1966. Descrip- of herring larvae 22-40 mm in length was higher on tion of embryonic and early larval stages of bluefish, Pomcu live Tigriopzcs than on pieces of squid flesh, which tomus salatrix. Trans. dm. Fish. Hoc., 95 : 264-271. indicates that even advanced larvae prefer live food. Fabre-Domergne and E. BiCtrix. 1897. Recherches biologiques npplicnhles ;i la piscimltnre maritime sur les axfs et les larves des noissons de mer et siir le tiirbot. Ann. Sci. Nat. ACKNOWLEDGMENTS (Zool.), 8 SCr., Tome 4: 151-220. -. 1902. Appareil ;i rotation pour 1'6levage des sufs de larves I wish to thank Lillian Vlymen of the Fishery- des poissons marins. C. R. Ass. Franc. Av. Sci. Ness. 30, Oceanography Center, National Marine Fisheries 1902, Pt. 2 : 577-582. -. 1905. Developpement de In Sole (&"lea vulgaris). Intro- Service, La Jolla, for translating papers from the duction R I'btiide de la pisciculture marine. Travail du Russian. and Dr. Peter Miyake of the Inter-American Laboratoire de Zoologie Maritime de Concarneau. Vnibert et Tropical Tuna Commission for translating portions Nony. Paris. 243 pp. of several Japanese papers. Fahey, W. E. 1964. A temperatnre controlled salt-water circu- lating apparatus for developing fish eggs and larvae. J. Cons. Int. Explor. Mer, 28 : 364-354. REFERENCES Fishelson, 11. 1963. Ohserrations on littoral fishes of Israel. 11. Larval development and metamorphosis of Blennius pavo Anthony, R. 1910. The cnltivation of the turbot. Proceedings Risso (Teleostei, Blenniidae). Israel J. Zool., 12: 81-91. of the 4th International Fishery Congress, Washington, 1908, Fliichter, J. 1965. Versuche ziir Brntaufzucht der Seezunge Pt. 2. Bull. U.S. Bur. Fixh. 28:861-870. (Translation.) Nolea solea in kleinen Aqiiarien. Helgol. zoiss. Meeresunters., Bardach, J. E. 1968. The status and potential of aquaculture, 12 : 395403. (English summary.) particularly fish culture. Vol. 2, Part 111. Fish culture. Forrester, C. R. 19M. Laboratory observations on embryonic Amer. Inst. Biol. Sci., Washington, D.C., 225 pp. (Pages development and larvae of the Pacific covl (Gadus macro- 22-29 of this work describe the rearing experiments of G. 0. cephalus Tilesins). J. Fish. Res. Bd. Canada, 21: 9-16. Schnmann.) Fnjita, S. 1957. On the larval stages of a scorpaenid fish, Se- Bishai, H. M. 1961. Rearing fish larvae. Bull. Zool. SOC.Egypt. bastes pachycephalus nigricans (Schmidt). Jap. J. Ichthyol., (16) : 4-29. 6: 91-93. (In Japanese with English summary.) Blaxter, J. H. S. 1962. Herring rearing-IV. Rearing beyond -. 1965. Early development and rearing of two common flat- the yolk-sac stage. Nar. Res. Scot., (1) : 18 pp. fishes, Eopsetta grigorjezvi (Herzenstein) and Tanakius kita- __ 3965. The feeding of herring larvae and their ecology in . harai (.Jordan et Starks). Bull. Jap. SOC. Sci Fish., 31: relation feeding. Calif. Coop. Oceanic Fish. 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Hirano, R., and Y. Oshima. 1963. On the rearing of larvae of Morris, R. W. 1955. Some considerations regarding the nutri- marine animals with special reference to their food organisms. tion of marine fish larvae. J. Cons. Int. Explor. Mer, 20: Bull. Jap. SOC.Sei. Fish., 29 : 282-297. (In Japanese.) 25.5-265. Ivanchenko, L. A., and 0. F. Ivanchenko. 1969. Transition to -. 1956. Some aspects of the problem of rearing marine active feeding by larval and juvenile white sea herring (Clu- fishes. Bull. Inst. Oceamogra. Uanaco, (1052) : 61 pp. pea pallosii Natio Maris-albi Berg) in artificial conditions. Nikitinskaya, I. V. 1958. On the onset of active feeding of the Dokl. Biol. Sei., 184 : 207-209. (Translated from Dokl. Akad. larvae of Clupea harengus pallasi. Val. 2001. Zh., 37: 1568 Nauk S.S.S.R., 184 : 1444-1446.) -1571. (In Russian with English summary.) Joseph, E. B., and V. P. Saksena. 1966. Determination of sa- Okamoto, R. 1969. Rearing of red sea bream larvae. Bull. 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Invest. Rept., 14 : fish. Preliminary communication. Ropp. et Proc.-Verb., Cons. 84-87. Int. Explor. Mer, 109, Pt. 3; 133. Kurata, H. 1956. On the rearing of larvae of the flatfish, Li- -. 1940. Ctklekking og oppdretting av saltvannfisk. Naturen, opsetta obscura, in small aquaria. Bull. Hokkaido Reg. Fish. (6/7) : 397-217. Res. Lab., 13 : 20-29. (In Japanese with English summary.) Rosenthal. H. 1969a. Verdauiingsgeschwindiglieit, Nahrungswahl __ . 1959. Preliminary report on the rearing of the herring iind Kahrungsbedarf Ijei den Larven des Herings, Clupea larvae. Bull. Hokkaido Reg. Fish. Res. Lab., 20 : 117-138. harengus 1,. Ber. Dt. Wiss. Iiomm. dfeeresforsch., 20: 60-69. (In Japanese with English summary.) (English summary.) Lasker, R., H. &!I.Feder, G. H. Theilacker and R. C. May. -. 196%. Untersuchungen iiber das Beutefangverhalten hei 1970. Feeding, growth and snrrival of Engraulis mordax Larven des Herings Clupea harengus 1,. Mor. 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Skr., Ser. Ilavunders@k.,G(8) : 71 pp. McMynn, R. G. and W. S. Hoar. 19233. Effects of salinity on Schach, H. 1939. Die kiinstliche Aafzucht van Clupea harengus the development of the Pacific herring. Canad. J. Zool., 31: L. Ifelgol. wiss. ilfeercsunters., 1 : 359-372. 417432. Shelhourne, a. E. 1964. The artificial propagation of marine fish. Meyer, H. A. 1878. Biologische Beobachtungen bei kunstlicher Ad!.. Mar. Biol., 2: 1-S3. Aufzucht des Herings der Westlichen Ostsee. Weigandt, Hem- Shojima, Y. 1957. On the de~elopnientof eggs and rearing of pel and Parey, Berlin. (Translation in Rep. U.S. Comm. larvae of a puffer, Fugu (Higanfugu) pardalis (T. and S.). Fish., 1878, Pt. 6 : 62W338.) Sei. Bull. Fac. Agr. Iiyushu Univ.,16: 125-136. (In Japa- Mito, S., M. Ukawa and AI. Higuchi. 1969. On the development nese with English summary.) and rearing of the larvae of a flonnder, Kareius biocoloratus Soleim, P. A. 1942. Arsaker til rike og fattige hganger av sild. (Basilewsky) with reference to its spawning in the culturing Fiskei-idir. Skr., Ser. HavundersBk., 7(2): 39 pp. (English pond. Bull. Nansei Reg. Fish. Res. Lab., 1: 87-102. (In summary. ) Japanese with English summary.) Tamamoto, G., and C. Kishioka. 1952. The development and Molander, A. R., and M. Molander-Swedmark. 1957. Esperi- rearing of hatched larvae of North Pacific Cod (Gadus mental investigations on variation in plaice (Pleuronectes nkacrocephalus Tilesius). Spec. Puhl. Jap. Sea Reg. Fish. platessa I,.), ltept. Inst. Nor. Res. Lysekil, Ser. Biol., (7) : Res. Lab., on the third anniversary of its founding: 301-308. 45 PP. (Translation no. 402, Fish. Res. Bd. Canada.)