Feeding Larval Marine Fishes in the Laboratory: a Review Robert C
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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” ____