ANNALES 13/'98 original scientific paper UDC 597.5(262.3-11) EARLY LIFE HISTORY STAGES OF THE FAMILY CARANCIDAE IN THE EASTERN ADRIATIC Jakov DULCiC Institute of Oceanography and Fisheries, HR-21G0G Split, P.O. BOX 500 ABSTRACT Use of early life history stages in systematic and ecological studies of fish has increased in recent years. It is now recognized that eggs and larvae present a wide array of characters that are largely independent of adult characters and, as such, suitable for a taxonomic analysis. On the other hand, fisheries recruitment studies focus on the survival of eggs and larvae as the mosi important factor influencing variations in population abundance. A requisite to these studies is detailed information on the appearance offish eggs and larvae in order to identify them in plankton sam­ples. Family Carangidae is of great interest to the Croatian fishery. This paper presents all available information and results on the early developmental stages of the family Carangidae found in the eastern Adriatic. Key words: early developmental stages, Carangidae, eastern Adriatic INTRODUCTION The importance of eariy-life-history studies to fisher­ies investigations and phylogenetic research has in­creased dramatically during the last decade. Earîy-life­history stages are now routinely used in fisheries studies to investigate the interannual variation in recruitment (Wooster, 1983), and in the studies of the phylogeny of fishes (Moser et al., 1984). The study of fish eggs and larvae is a key component in research into the biology, •systematic s and even population dynamics of fishes, in that it provides information on spawning areas and peri­ods of many species. By combining the location of the eggs, larvae and adults of a species with information on the surrounding environment, possible environmental effects on spawning (egg and larval transport, etc.) can be inferred. Studies of this type contribute to our under­standing of the early stages of development of fishes, which for certain species are still completely unknown. Consideration of the factors that affect egg and larval survival is fundamental, since it is the early stages of de­velopment that will eventually determine the existence of good or bad year classes. This is one of the main thrusts of ichthyoplankton studies in those areas, in which the requisite basic information is available (egg <3nd larval surveys in spawning areas during the spawn­ing season, etc.) for the species of interest. Family Carangidae is of great interest to the Croatian fishery, it is not possible to get new data on catch for every species now, but Grubiá!. (1982) reported that it is around 650 tons per year for Atlantic horse mackerel (Trachurus trachurus - more than 50%) and Medi­terranean horse mackerel (Trachurus mediterraneus ­less than 50%), 3 tons for pompano (Trachinotus ova­tus), and around 30 tons for yellow tail (Serióla dumer­ m. The aim of this paper is (1) to compile and present all the available information and results on both the early development stages of the eggs and larvae of the family Carangidae found in the eastern Adriatic and (2) to identify possible spawning areas and seasons of some species from the family. MATERIAL AND METHODS The present paper sets out descriptions of the eggs, yolk-sac larvae and larvae of the family Carangidae likely to be collected in plankton samples in the eastern Adriatic, together with information on the possible spawning areas and seasons for some species. The egg and larval descriptions have mostly been taken from the existing literature. In some cases the descriptions were done by the author himself based on material collected during surveys; in other instances the descriptions were 55 ANNALE S 13/'98 Jakov DUi.ČiČ: EAR t. V LIFE HISTORY STAGES OF THE FAMILY CARANGIDAE IN THE EASTERN ADRIATIC, .5S-64 f Fig, 1: Trachurus trachurus (L.). (a) egg; (b) yolk-sac larva, 2.6 mm; (c) yolk-sac larva, 3.2 mm; (d) larva, 4.9 mm; (e) larva, 5,2 mm; (f) larva, 6.7 mm (after Heincke and Fbrenbaum, 1900). Si. 1: Navadni šnjur Trachurus trachurus (L). (a) Ikra; (b) larva z rumenjakovo vrečko, 2,6 mm; (c) larva z rumenjakovo vrečko, 3,2 mm; (d) larva, 4,9 mm; (e) larva, 5,2 mm; (f) larva, 6,7 mm (po Heinckeju in Ebrenbaumu, 1900). !he North Sea {Heincke & Ehrenbaum, 1900). Holt (1893) obtained ovarian eggs from Trachurus trachurus brought in by mackerel boats on 16 June at Penzance. He thus established that the egg was pelagic and had an oil globule. Later (Holt, 1894) he obtained ripe unfertil­ized eggs from the North Sea on 19 May and described the segmented yolk. He also recorded that in some eggs there were two or three small oil globules which later coalesced into a single large one. When the embryo is half round the yolk brownish-yellow and black pigment appears along the body contours and especially round the oil globule. Ehrenbaum (1905-09) states that the black pigment appears first, and that the other colour is more brownish-yellow by reflected light but appears more yellow by transmitted light. Holt (1897) recorded differences in sizes of eggs from the North Sea and from Plymouth as follows: Grimsby - diameter of egg - 1.03­ 1.09 mm and diameter of oil globule - 0.26-0.27 mm; Plymouth - diameter of egg 0.81 -0.93 mm and diameter of oil globule - 0.22-0.23 mm. Polonskii and Tormosova (1969) made artificial fertilizations of eggs from parents 27-30 cm long. Twenty-four hours after fertilization the diameters of the eggs were 0.9-1.05 mm (mean 1.0 mm) and of the oil globules 0.25-0.275 mm (mean 0.269 mm). Kiliachencova (1970) reported next measures of eggs: diameter 0.90-1.10 mm and oil globule diameter 0.15-0.20 for the waters of Northwest Africa. Demir (1961) presented measure for Atlantic horse mackerel eggs from Sea of Marmara and the Black Sea: diameter is between 0.79 and 0.95 mm. An unscuiptured elastic eggshell protects a completely segmented yolk with usually one oil globule at its upper surface. The eggshell and yolk are transparent and colourless. The oil globule is a light copper colour in living material. There are melanophores on the oil globule and body of the em- published by other researchers, either for eggs and lar­vae actually collected in the eastern Adriatic itself or for egg and larval material collected in other areas but for species that also inhabit the waters of the eastern Adriatic. Notochord (NL) length was measured for pre-flexion, standard length (SL.) and total length (TL) for flexion larvae. RESULTS AND DISCUSSION Trachurus trachurus (1.) Eggs Pelagic, spherical, 0.81-1.04 mm in diameter, seg­mented yolk, oil globule 0.19-0.28 mm in diameter for Fig. 2: Trachurus trachurus (L.). Opercular spines of larvae: (a) and (b) 3.5 mm, (c) 5.0 mm, (d) and (e) 6.5 mm (after Schnakenbeck, 1931). SI. 2: Navadni šnjur Trachurus trachurus (L). Škržni trni larv: (a) in (b) 3,5 mm, (c) 5,0 mm, (d) in (e) 6,5 mm (po Schnakenbecku, 1931). 56 ANNALES 13/'98 jakov DUlOC : EARLY LIFE HISTORY STAGES OF THE FAMILY CARANGIDAF IN THE EASTERN ADRIATIC, 35-64 bryo after blastopore closure. The perivitellin space is narrow (Demir, 1961). Duidc (unpublished data) found in May at StonCica station (43°00'N 16°20,E - 4N m SE from island Vis in the middle Adriatic) horse mackerel eggs with the following measures: diameter 0.93-1.08 mm and oil globule diameter 0.17-0.22 mm (Fig. 1 a). Yoik-sac larvae DulCic? (1992) found Atlantic horse mackerel yolk-sac larvae at StonCica station in February, March, and April with a frequency 0.40%. The newly hatched yolk-sac larvae is about 2.5 mm long (Heincke & Ehrenbaum, 1900) (Fig. lb). The most characteristic feature is the anterior position of the oil globule in the segmented yolk and the extension of the anterior end of the yolk sac beyond the anterior margin of the head. The anus is situated behind the posterior end of the yolk-sac; as the larvae grow it reaches about the mid-point of the body. When newly hatched the pigmentation consist of melanophores and brownish-yellow chromatophores. The melanophores are at first irregularly distributed over the body, but as develop­ment proceeds the pattern gradually changes. At first the tail region becomes free of melanophores and a few melanophores appear on the dorsal and ventral pri­mordial fin on the level of the rectum, together with brownish chromatophores to form a distinct ventral bar. One or two marginal melanophores also appear on the anterior half of the dorsal fin and together with the brownish chromatophores tend to form a bar at the posterior end of the yolk-sac. The posterior half of the oil globule is well pigmented. By the time the yolk is nearly fully absorbed and the eyes are pigmented (length of 3.24 mm), the pigmentation is reduced to dor­sal and ventral body contour rows of melanophores, and a fringe of brownish pigment along the margins of the dorsal and ventral fin (Fig. 1c). Larvae }. Karlovac. (1967) found five Atlantic horse mackerel larvae at the Split channel station. O. Karlovac & J. Karlovac (1971) reported about 620 yoik-sac larvae and larvae found (length from 2.37 to 19.14 mm) mainly at the stations in the south and middle Adriatic during the expedition "Hvar" (1948-1949) from February to July. Regner (1980, 1982) observed larvae at the station in Kastela Bay (43-31'N 16°19'E) in February, April, and October with frequency 0,24%, and at the Stondca sta­tion from February to August with frequency 1.44%. In the earliest stages (3.5-4 mm) the pigmentation pattern consists of dorsal and ventral body contour rows of melanophores, with the beginnings of a mediolateral row. The dorsal row ends about mid-way between the anus and the caudal end and consists of about 10 large Fig. 3: Trachurus mediterraneus (Steindachner, 1868). (a) yolk-sac larva, 2.1 mm, (b) yoik-sac larva, 2.6 mm, (c) larva, 3,2 mm, (d) larva, 4.55 mm, (e) larva, 5.1 mm, (f) larva, 7 mm, (g) larva, 10 mm, (h) larva, 14 mm, (i) larva, 20.2 mm (after Demir, 1961). SI. 3: Sredozemski šnjur Trachurus mediterraneus (Steindachner, 1868). (a) larva z rumenjakovo vrečko, 2,1 mm, (b) larva z rumenjakovo vrečko, 2,6 mm, (c) larva, 3,2 mm, (d) larva, 4,55 mm, (e) larva, 5,1 mm, (f) larva, 7 mm, (g) larva, 10 mm, (h) larva, 14 mm, (i) larva, 20,2 mm (po Demirju, 1961). 57 ANNALES 13/'98 jnkov. DUIÜ Ó EARLY LIFE HISTORY STAGES O F THE FAMILY CARANCIOAE IN THE EASTERN ADRIATIC. S5-64 If Fig. 4: Pseudocararix dentex (Bloch & Schneider, 1801). (a) larva, 6.0 mm (after Schnakenbeck, 1931), (b) jaws, (c) opercular spines. SL 4: Vrsta Pseudocararix dentex (Bloch & Schneider, 1801). (a) larva, 6,0 mm (po Schnakenbecku, 1931), (b) čeljusti, (c) škržni trni, stellate meianophores opposite those of the dorsai row followed by a number of small meianophores extending nearly to the end of the urostyle. There are also numer­ous meianophores on the lower jaw, along the abdomen and the upper surface of the stomach, and on the head. As the larvae grow, the number of meianophores in­creases to cover gradually the sides of the body, leaving the caudal end free, except for the ventral row. The mediolateral row persists and the lateral body pigment extends forward. At a length of 6 mm a row of meiano­phores appears along the anal fin midway between the anus and the tail, situated along the outer margin of the interspinous area. The swimbladder is already apparent when the larvae is about 4 mm long. The urostyie begins to turn up at about 6 mm length. The interspinous areas of the dorsal and anal fins begins to develop at a length of about 5 mm, the dorsai about midway between the anus and the tail, and the anal at first anteriorly near the anus {Figs. I d and 1e). The fin rays of D^, D2 and A have developed by the time the larvae are 6-7 mm long and the fish is beginning to assume its carangid appear­ance. The pelvic fin is becoming evident at lengths of 7­8 mm (Fig. If). A noticabie feature of the larvae is the height of the anterior half of the body and of the head. The upper jaw has small denticles along its margin and there are opercular spines {Fig. 2); the latter are not noti­ cabie unless specially looked for. Details of their devel­opment have been described by Schnakenbeck {1931). In individuals of the same size there is some slight variation in form and size of the spines and in their numbers {Fig. 2). The basic arrangement is a row of longer stronger spines on the posterior margin of the preoperculum, with an inner row of smaller spines on the crista. The spine at the angle of the preoperculum is the longest. In larvae 5 mm long there are about 4-5 spines on the outer row, and 5 on the inner row. in gen­era! the larger larvae have the greatest number of spines. The spines are already apparent in larvae 4 mm in length. Trachurus mediterraneus (Steirtdachner, 1868) Eggs Pelagic, spherical, 0.71-0.90 mm in diameter, seg­mented yolk with usually one oil globule at its upper surface, perivitelline space narrow (Demir, !961). No eggs attributable to this species have been collected so far during the surveys carried out in the eastern Adriatic. Yolk-sac larvae Yolk-sac larva has segmented and oval yolk-sac. The anterior edge of the yolk-sac slightly passes beyond the anterior point of the head. The oil globule is located at the anterior part of the yolk-sac. The anus is welt behind the yolk-sac, and the part of the body anterior to the anus is longer than the part of body posterior to the anus {Fig. 3a). Demir {1961) supposed that the size of the yolk-sac of Trachurus trachurus should be larger than that of Trachurus mediterraneus. The most reliable characteristic which differentiates yolk-sac larvae of those two species ts the meianophore which develops on the primary dorsai fin in the yolk-sac larvae of Tr.trachurus, while none develop in yolk-sac larvae of Tr. mediterraneus. The larva! size at the end of the yo!k-sac stage of Tr. mediterraneus is smaller than that of Tr. trachurus (Demir, 1961). No yolk-sac larvae attributable to this species have been collected so far in the eastern Adriatic. Larvae All subsequent larval developmental stages {e.g. the first appeareances of jugai spines, fins, the upward incli­nation of the posterior part of urostyie etc.) are reached earlier by the larvae of Tr. mediterraneus than by the (arvae of Jr. trachurus. Besides, the meianophores on the larvae of Tr. mediterraneus are lesser in number but larger in size than the meianophores on the larvae of Tr. trachurus {Fig. 3b to Fig. 3i). However, the distinguish­ing characteristic of the I arvae of both spcciss are the ANNALES 13/'98 Fig. 5: Naucrates ductor (Linnaeus, 1758). (a) egg; (b) larva, 5.4 mm; (c) larva, 6.3 mm, (d) larva, 6.9 mm, (e) larva, 11.7 mm, (f) larva, 16.0 mm, (g) larva, 20.8 mm (after Sanzo, 1931). SI. 5: Pilot Naucrates ductor (Linnaeus, 1758). (a) ikra; (b) larva, 5,4 mm; (c) larva, 6,3 mm, (d) larva, 6,9 mm, (e) larva, 11,7 mm, (f) larva, 16,0 mm, (g) larva, 20,8 mm (po Sanzu, 1931). ANNALE S 13/'98 jakov DULft 6 EARLY LIFE HISTORY STAGES OF THE FAMILY CARANGIDAL IN THE EASTSRN ADRIATIC, 55-64 meianophores which develop on the preanai ventral primary fin, on the ventral side of the belly posterior to ventral fin, and on the jaws in the larvae of Tr. trachu­rus. Corresponding chromatophores are either never found or are unimportant in number and size in the lar­vae of Tr. rnediterraneus (Demir, 1961). Larvae of this species were found in the southern Adriatic by Karlovac (unpublished data), and at StonCica station (June; n=2, 5.1 and 5.2 mm SL) by DulCic; (unpublisheddata). Pseudocaranx dentex (Bfocb & Schneider, 1801) Eggs No information available. Larvae Regner (1982) found gueliy jack Pseudocaranx den­tex larva at Stondca station in August with frequency 0.02%. Very little information exists concerning the larva! development of this species; only descriptions of two larvae measuring 5 and 6 mm, respectively, by Schnakenbeck (1931) and Padoa (1956) are at hand. The teeth develop early. No supraoccipital crest is pre­sent. The preopercular spines are distinct from those of horse mackerel larvae; this is most apparent on the up­per anterior margin of the preopercule which, unlike that, on horse mackerel larvae, is devoid of spines. The most outstanding feature of the pigmentation pattern is the nearly complete lack of meianophores along the body {Fig. 4). Naucrates ductor {Linnaeus, 1758) Eggs The eggs are spherical, 1.32 mm in diameter, with an oil globule 0.28 mm in diameter. The yolk is seg­mented and the perivitelline space is very narrow (Fig 5a). The chorion is smooth (Sanzo, 1931). No eggs at­tributable to this species has been collected in the east­ern Adriatic. Yolk-sac larvae No information available. Fig. 6: Serióla dumerili (Risso, 1810). (a) egg, (b) yolk-Larvae sac larva, 3.60 mm, (c) yolk-sac larva, 4.40 mm, (d) larva, 4.80 mm, (e) larva, 9.72 mm (after Sanzo, 1933). DulCiC (unpublished data) found pilotfish larvae at SI. 6: Gof Serióla dumerili (Risso, 1810). (a) ikra, (b) Stoneica station in August (n-2, 5.6 and 7.0 mm SL, re- larva z rumenjakovo vrečko, 3,6 mm, (c) larva z ru-spectively). menjakovo vrečko, 4,4 mm, (d) larva, 4,8 mm, (e) The head is very prominent, representing 32% of NL larva, 9,72 mm (po Sanzu, 1933). in very young larvae and 37% in 7 mm larvae. The 60 ANNALE S 13/9 8 Jakov DtJIČIČ: EARLY LIFE HISTORY STAGES OF THE FAMILY CARANGIOAE IN THE EASTERN ADRIATIC, 55-64 Fig. 7: Trachinotus ovatus (Linnaeus, 1758). (a) larva, 2.52 mm, (h) larva, 5.36 mm, (c) latva, 5.56 mm (after De Gaetani, 1940% (d) juvenile, 73.8 mm (after Dulčič et al., 1997). Si 7: Vrsta Trachinotus ovatus (Linnaeus, 1758). (a) larva, 2,52 mm, (b) larva, 5,36 mm, (c) larva, 5,56 mm (po De Caetaniju, 1940), (d) mladostni primerek (po Dulčiču etai, 1997). shape of the body subsequently changes, with the rela-to a certain number of very dark, transverse stripes alter­tive head depth decreasing as body depth increases, the nating with lighter areas. Notochordal flexion takes two approaching each other. The mouth is very small. place at between 5.5 and 6.5 mm SL (Figs. 5b and c). Long preopercular spines, supraorbital crest, and two The upper rays of the pectoral fins are discernible in postemporai spines are present on the head from early even the smallest larvae. These fins grows larger in later stages of development. The body is heavily pigmented stages, reaching the level of the vent in 7 mm SL larvae. from the earliest stages of development, except on the Ventral fin buds are also visible in very small larvae and caudal peduncle. The pigmentation pattern begins to are located at the level of the base of the pectoral fin. change at a length of 7 mm (Fig. 5d); at the beginning The final number of dorsal and anal fin rays is attained the black pigmentation extends onto the fins; later the at a larval length of more than 7 mm SL (Figs. 5e, f and pigmentation of the body becomes uneven, giving rise g) (Sanzo, 1931). 61 ANNALES 13/98 jskov DULCI Ó EARLY LITE HISTORY STAGES OF THE FAMILY CARANC1DAE IN THE EASTERN ADRIATIC, 55-61 Serióla dumeriU (Risso, 1810) Eggs Pelagic, spherical, transparent, with a large colour­less oil globule and slightly streaked capsule, segmented yolk, 1.04-1.12 mm in diameter, oil globule 0.28 mm in diameter (Fig. 6a) (Sanzo, 1933). Thirty hours after ferti­lization the embryo encloses 2/3 of the meridian and contains about 20 segments. Black and yellow pigmen­tation is present on the cephalic region and on the entire trunk. Stellate melanophores are obviously present on the yolk-sac (Sanzo, 1933). No eggs attributable to this species have been collected during any of the survey carried out in the eastern Adriatic. Yolk-sac larvae The newly hatched yolk-sac larvae is about 3.5 mm long (Fig. 6b) (Sanzo, 1933). The most characteristic feature is, as in horse mackerel larvae, the anterior posi­tion of the oil globule in the segmented yolk, which ends a notable distance from the anal opening. The mouth is not yet open. The anal opening is slightly be­hind the middle of the body. Melanophores are irregu­larly distributed on the body and some of them are on the oil globule (Fig. 6b) (Sanzo, 1933). No yolk-sac lar­vae attributable to this species have been collected during the surveys so far carried out in the eastern Adriatic. Larvae O n the third day after hatching the length increases to 4.40 mm, the mouth is open and the eyes depig­mented. Some melanophores are along the dorsal fin, on the ventral side of the body, and around the middle of the caudal region (Fig. 6c). When larva reaches 4.70 mm TL, the body is rather rough with a large head, the length of which is, in respect to the body, equal to 1/3 of the total length (Fig. 6d). Four short spines are in front of the preoperculum and three very long ones at the mar­gin behind the preoperculum (Fig. 6e). Stellate melano­phores are abundant all over the body except on the last part of the caudal fin, which is slightly more transparent. In larva about 9.72 mm TL, the body has suffered a no­table lateral compression and the preoperculum with spines has been reduced. The abdominal trunk and caudal are uniformly intensively pigmented except the peduncle caudal. In the cephalic region the pigment is thicker on the dorsal side (Fig. 6e) (Sanzo, 1933). Regner (1982) found 5. dumerili larvae at Stoncica station in July with frequency 0.02%. Trachinotus ovatus (Linnaeus, 1758) Eggs No information available. Yolk-sac larvae No information available. Larvae De Gaetani (1940) described the larvae in the stage of 2.52 mm in total length; body is slender and head well developed. Eyes are slightly oval and large in pro­portion to the head. An intensive pigmentation is present on the entire abdominaf region and on the dorsal on the head, and on the ventral and lateral region (Fig. 7a). At the stage 4.80 mm, orbital crest is notable on the ce­phalic region. The pigmentation was not modified, but the melanophores on the preoperculum and along the mandibular margin have become more numerous. In larvae about 5.36 mm (Fig,. 7b) the pigmentation has modified slightly and the preoperculum is scarcely more developed. In respect to the previous stage the orbital crest, on the contrary, is much more evident with 3-4 spines (Fig. 7c). There is a large chromatophore in the space between the third and fourth dorsal rays. Juvenile juvenile specimen of pompano T. ovatus was caught near sandy beach "Zlatni rat" in the vicinity of a small island settlement Bo! (island Brae) in December 1994 (Outfit ei a/., 1997). This specimen has small teeth in both jaws, consisting of a narrow band anteriorly, taper­ing posteriorly (Fig. 7d). Bases of soft dorsal and anal fins are approximately equal in length; pectoral fins are longer than pelvic fins. Spinous dorsal fin has 6 spines. There are no caudal peduncle grooves. Lateral line is slightly arched above pectoral fins and straight thereaf­ter. There are no scutes. The colour of the specimen is dark green on the top, silvery below. There are 4 dark vertically elongated blotches on the sides; distal half of dorsal and ana! fin lobes and tips of caudal fin are black. ANNALES l3/'98 Jakov DULČlC: EARLY LIFE HISTORY STAGES OF THE FAMILY CARANGIDAE IN THE EASTERN ADRIATIC, 55-64 ZGODNJ I RAZVOJN I STADIJ I RIB I Z DRUŽIN E CARANGIDA E V VZHODNE M JADRAN U Jakov DULČIČ Institut za oceanografiju i ribarstvo, HR-21000 Split, P.P. 50 0 POVZETEK Pomen zgodnjih razvojnih stadijev v sistematičnem in ekološkem raziskovanju rib postaja v zadnjih ietih vse večji. Tako je danes že splošno uveljavljeno prepričanje, da ikre in larve pomenijo širok spekter bitij, ki so v veliki meri neodvisna od odraslih bitij in kot taka primerna za taksonomske analize. Po drugi strani pa se študije o prirastku ribjih populacij osredotočajo na preživetje iker in larv kot najpomembnejšega dejavnika, ki vpliva na spremembe v številnosti ribjih populacij. Prvi pogoj za te študije so podrobne informacije o videzu iker in larv, da jih lahko identificiramo v planktonskih vzorcih. Najpomembnejša za hrvaško ribiško je družina Carangidae. V tem članku so zajeti vsi razpoložljivi podatki in rezultati o zgodnjih razvojnih stadijih rib iz družine Carangidae v vzhodnem Jadranu. Ključn e besede: zgodnji razvojni stadiji rib, Carangidae, vzhodni jadran LITERATURE Demir , M. 1961 . O n the eggs and larvae of the Trachutvs trachurus (L.) and Trachurus mediterraneus (Stahnr.) from the Sea of Marmara and the Black Sea. Rapp. P. -v. Reun. Cons, int. Explor. scient. Mer Medit., 16 (2): 31 7-320. De Gaetani, D. 1940. Stadii larvali e giovanili di Lichia glauca Risso, Mem. R. Com. Talassogr. ItaL: 560-564. Dulcid, J, 1992 . Istra2ivanje sastava i brojnosti ihtio­pfanktona u srednjem jadranu. Morsko ribarstvo, 3: 73­78, Dulcit,J,, A. Pailaoro & M. Kraljcvic, 1997. First record of pom pa no finger! in g, Trachinotus ova tu s (Linnaeus, 1758) (Pisces: Carangidae), in the eastern middle Adriatic. Nat. Croat. 6 (1): 61-65. Ehrenbaum, E. 1905-09. Eier und larven von Fischen der Deutschen Bucht. III. Fische mit fsetsitrenden. Giern Wiss. Meeresunt, Helgoland, vol. V!: 127 p. Grubisid, F. 1982. Ribe, rakovi i Skoljke Jadrana. STRO Naprijed, Zagreb-GRO Liburnija, Rijeka: 240 pp. Heincke, F. & E. Ehrenbaum 1900. Eier und Larven von Fischender deutschen Bucht. Ii. Die Bestimmung der schwimmenden Fischeier und die Methodik der Eim­pssungen. Wiss. Meeres untersuch. Helgoland, N.F., 3: 127-332. Holt, E. W . L . 1893 . Survey of fishing grounds, west coast of Ireland, 1890-91: on the eggs and larval and post-larval stages of teleosteans. Scient. Trans. R. Dublin Soc., Ser. 2, 5: 121 pp. Holt , E. W . L. 1894 . North Sea investigations. ]. mar. biol. Ass. U.K., 3: 169-201. Holt, E. W . L. 1897 . Preliminary notes on the reproduc­tion of teleostean fishes in the south-western district. ). mar. biol. Ass. U.K., 5: 41-50. Karlovac , J. 1967 . Etude de I'ecologie de la sardine, Sardina pilchardus Walb., dans la phase planctonique de sa vie en Adriatique moyenne. Acta Adriat., 2: 1-101. Karlovac, J. & O. Karlovac 1971. Contribution a la con­naissance de I'ecologie de Trachurus trachurus (L.) au large de I'Adratique. IzvjeSCa-Reports Institut za oceano­grafiju i ribarstvo, Split, V, 2A: 3-24. Kiltachenkova, V. A. 1970. Development and distribu­tion of eggs and larvae of Trachurus trachurus L. Rapp. P. -v, Reun. Cons. perm. int. Explor. Mer Medit., 159: 194-198, Moser, H. G., W. J. Richards, D. M. Cohen, M. P. Fahey, A. W. jr. Kendall & S. L. Richardson (editors), 1984 . Ontogeny and systematics of fishes. Spec. Publ. 1, Am. Soc, Ichthyol. Herpetol. Allen Press, Lawrence KS: 760 p. Padoa, E. 1956. Heterosomata. Fauna Flora Golfo Na­poli, Monogr., 38; 783-877. Polonskii, A. S. & I. D. Tormosova, 1969. The spawning of the horse mackerel of the north-east Atlantic and the distribution of its eggs and larvae (English translation). Trudy Atlant. Nauch. issl. Inst. Ryb. Khoz., 23: 27-48. Regner, S. 1980 . The larval stages of fish in Kastela Bay. Acta Adriat,, 21 (2): 123-136. 63 ANNALES 13/'98 Jakov DULClC: EARLY LIFE HISTORY STAGES OF THE FAMILY CARANGIDAE IN THE EASTERN ADRIATIC, 55-64 Regner, S. 1982. Istraživanja sastava i brojnosti larvalnib stadija riba u planktonu otvorenog mora srednjeg Jadrana. Studia Marina, 11-12: 45-60. Sartzo, L. 1931. Uova, stadi embrionali e post-em­brionaii di Naucrates ductor L. Me m. R. Com. Taiassogr, Ital., 185 pp. Sanzo, L. 1933. Uova e larve d i Seriola dumerili Risso. Mem. R. Com. Taiassogr. Ital.: 555-559. Schnakenbeck, W . 1931. Carangidae. Rep. Danish Oce­anogr. Exped. 1908-10 Medit adjac. Seas. 2, Biology, A. 14: 20 pp. Wooster, W. S. 1983. From year to year: International variability of the environment and fisheries of the Gulf of Alaska and the eastern Bering Sea. Rep. WSG-WO 83-3, Wash. Sea Grant Prog., Univ. Wash., Seattle, NA 98195: 208 pp.