INVESTIGATIONS OF FOOD AND FEEDING HABITS OF LARVAE AND JUVENILESOF THE ARGENTINE ANCHOVY ENGRAULIS ANCHOITA'JANINA DZ. DE ClECHOMSKllnstituto de Biologia MarinaMar del Plata, ArgentinaINTRODUCTIONUp to the present time no papers on the feeding ofthe larvae of marine fishes have been produced inArgentina. The present contribution is the first studyof this nature referring to this area of the SouthwestAtlantic and it deals with problems related to thefeeding habits of the larvae and juveniles of theArgentine anchovy.The purpose of this paper is to study the food andfeeding habits of the anchovy larvae from the time atwhich the larvae begin to be able to feed, to the timewhen the juveniles approach the adult stage. Thisstudy has been based upon a quantitative and qualitativedetermination of the food components, to obtaina better assessment of the contribution of the differentgroups of organisms to the diet of the larva(. andjuveniles of the anchovy. The characteristics of thefeeding of the young forms of the anchovy at differentstages of development were studied in relation to themorphological changes ~vhicli take place throughoutthe growth of the individuals. The morphologlcalcharacters most closely related to the feeding processin fishes are: the digestire tract, the gill rakers andthe dimensions of the mouth. Therefore, in the presentwork, special emphasis has been given to a detailedstudy of these morphological elements. Also, an analysisof the changes in the feeding of the larvae andjuveniles of the anchovy during different seasons ofthe year has been attempted.Together with the anchovies, juvenile indivudals ofanother species, Aicstroatlzerina incisn, were gathered.The individuals of this species share the habitat of,and are found mixed with those of the anchory. Thepurpose of collecting other material was to study theintestinal content of the individuals of this otherspecies with the aim of obtaining comparative materialas a frame of reference for the anchovy, taking especiallyinto account the competition for their foodsupply. The results of the analysis of the feedinghabits of the juveniles of A. incisa will be found inanother paper by this author (Ciechomski, in press).MATERIALS AND METHODSThe larvae and juveniles of anchovy which wereused as material for this study were collected fromcoastal maters off Mar del Plata. The juveniles wereobtained from every month of the year. The larvae ofsizes 3.0-4.9 mm and 5.0-22.0 mm were obtained onlylThis paper has heen iireparerl through the sponsorship of theConselo Wncioiinl 17s Invrst?gncio?les CientafLcns y Tecwwas ofAvr/eiittua.during the summer. A total of 1,705 larvae andjuveniles has been studied, ranging in size from3.0 to 90.0 mm. From these, 503 individuals containedfood, the remainder having their digestive tractsempty.For the elaboration and presentation of the data, themonths of September, October, and November havebeen considered spring, December, January, and Februarywere considered as summer, March, April, andMay autumn, and June, July, and August winter.The material to be studied was fixed on the spot,right after its collection, with formaldehyde, in orderto stop the digestive processes, since this was convenientin view of the study made of the intestinalcontent. Further treatment of the material was continuedin the laboratory.Each individual was measured and its digestivetract was separated under a magnifying glass. Theexamination of the intestinal content was performedunder the magnifying glass and the microscope. Allof the components vere separated, counted and measuredby means of a micrometric eyepiece. For assessingthe weight of the individuals, a list of meanweights was made in which the larvae and juvenilesof anchovy were classed in total length classes with2 mm intervals.Plankton samples were obtained together with theanchovies, and the predominant forms were noted.Although this appraisal was not made in a quantitativemanner and was not expressed numerically, it wasconsidered useful as an indication of the feedingselectivity of the fishes which were studied.In the quantitative treatment of the data, one ofthe methods used was the determination of the frequencyof occurrence. The calculation of the weightof the total food ingested and of the percentage compositionof the various food components was basedon the mean live weight of the planktonic organismswhich make up the diet of the anchovies. Since atthe present time no data are available on the weightof these planktonic organisms, which are the basicfood supply of the larvae and juveniles and, in somecases, of adult planktofagous fishes, for the most ofthe planktonic components the volume was determinedby the method of volume shift. In other cases, whendealing with very small organisms of irregular shape,such as some diatoms (Coscinodiscus, Triceratizm,etc.) the volume determination was accomplishedgeometrically. Multiplying the volume by the correspondinggroup index (Hagnieier, 1961) the meanweights of the organisms were obtained. The following
~table shows the values calculated for the volume andweight of some of the planktonic components.REPORTS VOLUAIE XI, 1 JULY 1963 TO 30 JUNE 1966 73TABLE 1WEIGHT OF SOME PLANKTONIC ORGANISMS MOST FREQUENTLYFOUND IN THE FOOD OF THE LARVAE AND JUVENILES OF1MARINE FISHESMean Hagmeier MeanOrganisms di:F volume coefficient weight(mm3)(mg)PeridineaEzuuiaello sp..----CopepodaCalanoida- ._._._. 1000" .~._. _._ 1000-2000" ........ 2000-3000" _._...__ 3000-4000" .~. 4000-5000-.. -Harpacticoida(Euterpina acnt.).Cyclopoida---. .DiatomeaCoseinodiseus spp.- 124 x 32Triceratium spp.. . 145 x <strong>11</strong>4 x 6(CladoceraEuadne nard .Larvae of Lamellibranch...........Anchovy eggs.. . . ..Fish eggs.. . . . . ~48 x 36480 x 850-._.356 x 176f240 x 130. 420-780200-3441370 x 72010000.0001600.0004960.00003250.01200.13300.45801.42804.00000.00950.00500.00920.00440.37800.52301.<strong>11</strong>.<strong>11</strong>.051.041.041.041.041.041.041.04(1.04)0.0001750.0005450.0000340.01250.13730.47141.48514.16000.01000.00520.0095..._ 0.0044_.__ 0.3780__.. 0.5230___Of course, these calculations are not free from error,especially in the case of the Copepoda, which havebeen classed by great groups and by size classes,instead of by separate species, which would havebeen more correct. In the present conditions, a morerefined treatment was not feasible.For the calculation of the ingestion coefficient thefood weight was divided by the weight of the individual,and the result was multiplied by 1,000.The measurements of the dimensions of the mouthwere made by means of the inicrometric eyepiece. Themouth was opened by means of very thin needles andthe length and width of the mouth were measuredunder approximately the same angle. The first branchialarch of the left side was used always for observingthe development of the gill-rakers.INCIDENCE OF FEEDINGThe incidence of feeding was remarkably low,especially in some of the size classes of larvae. InTable 2 the values for the incidence of feeding forlarvae of different lengths are shown.As is shown in Table 2, the incidence of feedingvaries greatly with the size of the larvae and juvenilesof the anchovy. It is relatively high for the larvaewhose lengths fall within the 3.0-4.01 mm class. Thelarvae in this group have just reabsorbed their yolkand have begun to feed themselves. While the growthof the larvae continues the incidence of feeding de-Length ofOrganism (mni)Number ofOrganismsIncidence ofFeeding3.0- 4.0 -...--..... 5652.24.0- 5.0 ._......... 4218.25.0- 9.0 .--.--..... 5009.0-20.0 ........... 1 3505.720.0-30.0 ..-.._..... 360 9.230.0-40.0 ........-..17.940.0-50.0 --..-...-.. I 57.350.0-90.0 ...........245 78.6Total ...........1705 larvae503-NumberIof organisms containing foodcreases sharply, reaching the value 0 for individuals inthe 5.0-9.0 mm class length. In those larvae whoselengths exceed 9.0 mm the incidence of feeding beginsto increase gradually, but at a low rate, until the timeat which the larvae reach a length of 40 mm. Beyondthe 40 mm length the number of larvae containing foodincreases clearly and in the juveniles of 50.0-90.0 mmthis number reaches a more or less constant and ratherhigh level. It should be emphasized that the fact thatlarvae which did not contain food were found in sucha scarce number has been experienced during 3 years.regardless of the season of the year.This low incidence of feeding found in larvae of aspecies of the family Engraulidae is not an isolatedfact, but has been observed with different species ofthis family and of those of the Clupeidae. Berner(1959) while studying the feeding habits of the northernanchovy, Engradis waordax found that amongthe 13,620 larvae which he examined only 2<strong>11</strong> had ingestedany food. Lebour (1921), who studied the foodhabits of numerous young clupeids, found that thepercentage of individuals containing food was verylow. The incidence of feeding in the larvae of othergroups of fishes appears to be higher (Lebour 1920,Wiborg 1948, etc). While studying the food of thelarvae and juveniles of A. incisa which were collectedtogether with those of the anchovy (Ciechomski, inpress) the present author found that all of the individualswhich were examined contained food in theirintestinal tract.The phenomenon of the low incidence of feeding inthe families Engraulidae and Clupeidae has been asubject of detailed analysis by numerous authors.There exist, as a consequence, several theories whichattempt to explain this fact. Some authors try to explainthis phenomenon as a consequence of the selectivityof the nets, which would tend to collect thoselarvae which were weak and underfed (Berner 1959,Soleim 1942, and others). Other authors think ofthis phenomenon as a consequence of the characteristicsof the digestion of these species, which, accordingto them, is very rapid. The old theory of Putter, accordingto whom fishes are able to feed upon theorganic matter which is dissolved in the water, hasbeen sustained by Morris (1955). This author thinksthis possible on the basis of the fact that the larvae
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