monas sp. ) was maintained at cell/ ml. In the 7th day after hatched out, the ration of rotifer was increased to individuals/ ml.

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1 50 (1) :91-102, 2004 A cta Zoologica S inica 3 33,, , mm,, , 100, mm , 37 h, 30 min mm 1, 115, 2, 3,, 18, 25, 27, 29,,, 57, 79,, 3 :,,, ;,, [ 50 (1) : , 2004 ] Morphological and developmental characters at the early stages of Cynoglossus semilaevis 3 the tonguef ish WAN Rui2Jing 33, J IAN G Yan2Wei, ZHUAN G Zhi2Meng Key Laboratory for Sustainable Utilization of Marine Fishery Resources, Ministry of Agriculture, Yellow Sea Fisheries Research Institute, Chinese Academy of Fishery Sciences, Qingdao , Shandong, China Abstract Based on the data and the samples from both the hatchery experiments carried out during 1987 to 1989 and from the large2scale seedling2rearing performed in the Laizhou Mingbo Fish Farm in the autumn of 2002, we observed and described the morphological and developmental characters in the early stages of the tonguefish Cynoglossus semilaevis G nther, In each late August during the experiment years, a pair of 135 hp trawlers was used to catch adult tonguefish in the central and western parts of the Laizhou Bay, the Bohai Sea. Normally, those adult fish with the gonad developing to stage were chosen to be the parent candidates. As chosen, the body size of the female fish was mm in standard length with a body weight ranged from to g while the body length and weight of the male fish were mm and g, respectively. The parent fish were transported to the hatchery and cultivated in the concrete pond with a volume of 50 m 3. The cultivation density was individuals/ m 2. The female and male pro2 portion was kept in 1 3. The cultivation pond was shaded by black curtain. During cultivation, the daily exchange rate of seawater was about 100 % or more by supplying re2circulation water and aeration was discontinuously given to maintain water quality. The parent fish were fed twice a day and rationed 3 % - 5 % of fish body weight. The priority categories of diets were clamworm, squill and clam meat. When the gonad of the parent fish matured after one2month culture to the middle or late September, artificial insemination could be performed to obtain fertilized eggs. The fertilized eggs were placed in a glass tank with a volume of 120 L to incubate. On incubating, the gentle re2circulation water and fresh seawa2 ter were supplied. The newly hatched larvae were cultured in the same glass tank by supplying gentle re2circulation of wa2 ter with gentle aeration. In the second day after hatching, the larvae were fed with rotifer B rachionus plicatilis twice a day and rationed at a food density of 5-10 individuals/ ml. The density of unicellular alga ( Chaetoceros m uelleri, Isochrysis galbana, Platy , ( ) 863 (2002AA603011) [ This research was funded by the grants from the Na2 tional Natural Science Foundation of China (No ) and the National 863 Plan of China (No. 2002AA603011) ] 33 (Corresponding author). ν 2004 Acta Zoologica Sinica E2mail : ac. cn

2 monas sp. ) was maintained at cell/ ml. In the 7th day after hatched out, the ration of rotifer was increased to individuals/ ml. In the 12th day after hatched out, the larvae were supplied with both rotifers (10-15 individuals/ ml) and copepod larvae (2-3 individuals/ ml), while, the density of unicellular alga was maintained at or above cell/ ml. In the 19th day after hatching, the juveniles were moved to an indoor concrete pond with a volume of 810 m 3 for further cultivation. Using a preheated/ auto2flow device, the daily water exchange rate was kept at 1/ 2 to 2/ 3 of the total culture volume. The pond bottom was punctually cleaned. Aeration was supplied 4 times a day, lasting 30 minutes each time. The juveniles were fed thrice a day with the food categories of either rotifer + copepod larva or rotifer + cope2 pod larva + A rtemia nauplii and rotifer + copepod larva + A rtemia nauplii + minced fish meat (particle size : mm). Normally, the rations of rotifer, copepod and A rtemia nauplii were maintained in the densities of 5 or 6 in2 dividuals/ ml, 3-6 and 3-5 individuals/ ml, respectively. But, the ration of minced fish meat was adjusted according to the feeding conditions. The cultivation temperature was kept at a range of and the p H value at Dissolved oxygen was maintained at a range of mg/ ml. A luminance about 500 lux was controlled. The samples of fish eggs, larvae, postlarvae, juveniles and fingerlings were collected and preserved in 5 % neutral formalin for observation and description of the morphological and developmental characters at the early life stages. The egg of the tonguefish was separate, spherical and buoyant with a diameter range of mm. The chori2 on was thin, elastic, lubricous and transparent. The egg was rich in oil globules, the numbers of which were variable from 97 to 125, about 100 in the majority. The diameter of oil globule ranged mm. Under the culture tempera2 ture of , the eggs began to hatch in 37 hours after fertilization and larvae were completely hatched out within 30 min. The body length of the newly hatched larva ranged from 2156 mm to 2168 mm in total length. 12day2old larva appeared pectoral fin bud. 1152day2old larva developed a cruising swimming mode. 22day2old larva presented the crown2like larval fin and started its exogenous feeding. Swim bladder occurred in 32day2old larva. Hereafter, the ontogeny embarked on the postlarval stages. The ontogeny of 182day2old larvae entered the juvenile stages henceforth. The right eye of 252day2old juvenile started to migrate upwards. The right eye of 272day2old juvenile migrated to vertex. Growing to 292day2old, the right eye of the juvenile completely migrated to the left side and the crown2like larval fin disappeared. The pectoral fin became entirely degenerate. Fin ray of each fin completely developed. The ontogeny of 572day2old juvenile went into the fingerling stages. The swim bladder of 792day2old fingerlings degenerated and their scales entirely devel2 oped. As observed, besides strict control on water temperature and quality, it is critically important to supply larvae with suitable diets soon after the yolk and oil globules are consumed away and larvae start to feed. Furthermore, postlarvae should be supplied with live or fresh diets, which is one of the vital elements of the technical components in terms of rear2 ing postlarvae at metamorphosis stages [ Acta Zoologica Sinica 50 (1) : , 2004 ]. Key words Tonguefish, Cynoglossus semilaevis, Larva, J uvenile, Fingerling ( Cynoglossus semilaevis G nther, 1873 ) ( Pleuronectiformes) ;, (1986) (Cynoglossidae) ( Cy noglossus Buchanan2 Hamiltou, 1822), 3,, lenciennes, 1844) ,, ;,, (1983) (Wakasa Bay), ( Cynoglossus joyneri G nther, 1878, 1995) ; Balakrishnan and De2 523 mm, 800 mm, vi. 280 mm (, 1988a),, Cynoglossus lida ( Bleeker, 1852 ),, Cynoglossus brevis,, 217, cynoglossus Hamilton2Buchanan, 1822 (, 1988b), (the Bay of Bengal), 72 ( Cynoglossus monopus (Bleeker, 1849), 1995), ; Ahlstrom et al. (1984) (1957) (Moji), 13 Cynoglossus robust us G nther, 1873 Cynoglossus abbreviat us ( Gray, 1832) ( Hypophthal michthys molit rix Cuvier et Va2 (1974) Cochin Backwater Cynoglossus puncticeps Richardson, 1846 G nther, 1862 Cynoglossus Cynoglossus arel ( Schneider, 1801) ; ; (1965) (1983) ( Shimabara ) ; ( 1988a, 1988b,

3 1 : 1993), 1/ 2-2/ 3,, 4, 30 min 3 / ( mm) 5, ind. / ml, 3-6 ind. / ml, mg/ L, 500 Lux , 135HP, 211, 50 m 3, / m 2, 1 3, mm ( mm, n = 20) 670 mm, g, 270,, mm, g, (103 7, n =,, 20), 100,, 100 %,,, , 0111 mm ( mm, n = 20) 3 % - 5 % , 9,, 15 min, L, 415 h, 515 h, 15 h,,, 17 h, 2015 h ,, 37 h, 30 min, 37 h, 120 L 212, mm (2163 1/ 4-1/ mm, n = 12) mm ( 2152, 1 h 5-10 min mm, n = 12) 0148 mm, , p H , mg/ L 2,, 0179 mm 40 2, 5-10 ind. / ml,,, ( Chaetoceros m uelleri, Isochrysis galbana, Platymonas sp. ) cells/ ml 7, ind. / ml, 12,, 1/ , (Copepod larvae) 2-3 ind. / ml,, cells/ ml 5 ind. / ml, 21-8, p H , , min, 115 h, 315 h ( 1 : 1-9) mm, 115,, 1114 mm,, 1124 mm ( 2 : 1) h,, 5 19, 810 ( 2 : 2 ; 3) m h, 93

4 , 1/ , 81 3/ 5 91 Fig. 1 Eggs of the tonguef ish 1. Fertilized egg cell stage. 3. Multi2cell stage. 41 Late blastula stage. 5. Middle gastrula stage. 6. Blastopore nearly closed. 7. Blastopore closed, Kupffer s vesicle formation. 8. Embryo encircling 3/ 5 of the yolk2sac. 9. immediately before hatching mm ( mm, n = 8), mm ( mm, n = 8), mm ( mm, n = 8),, ( 2 : 5),,,, 21 h, , mm ( mm, n = 5), mm, 5124 mm mm ( mm, n = 5),, mm ( mm, n = 5) 2-4,,,, 115 ( 2 : 3) 2 ( 1 ), mm ( mm, n = 8), mm ( mm, n = 8), 213,, 5 5 ( 2 : 6),, , ,, 5156 mm ( mm, n = 8), ( 2 : 4) 115, 5149 mm ( mm, n = 8)

5 1 : 2 11, 2156 mm 2. 6 h h,, 4160 mm 41 1, 5103 mm , 5141 mm Fig. 2 Prelarvae of the tonguef ish 1. Newly hatched larva, 2156 mm in total length hour2 old prelarva after hatching hour2old prelarva, the premodium of the crown2like larval fin appeared, 4160 mm in total length day2old prelarva, 5103 mm in total length day2old prelar2 va day2old prelarva, 5103 mm in total length., (118) (91),, 0128 mm,,,,, , 0126 mm,,,, (, 5 ) ( 5 : 1), , mm ( mm, n = 10),,, mm ( mm, n = ( 4 : 1) , mm, mm ( mm, n = 8), 0180 mm, 0134 mm, 5 ( 4 : 2) 5, mm ( mm, n = 8), mm ( mm, n = 8) 1110 mm 2 ( 4 : 3) 6, 6128 mm, 5188 mm, 1148 mm 0140 mm,,,,, ( 4 : 4) 8, 7110 mm, 6190 mm 1180 mm, , mm ( mm, n = 6), mm ( mm, n = 6) 2110 mm,, 2,, ( 4 : 5) , mm ( mm, n = 8), mm ( mm, n = 8), mm ( mm, n = 8) 16 10), mm ( n = 10), mm ( mm, n = 8), mm, 95

6 h Fig. 3 Dorsal view of 62hour2old larva, showing sensory tubes , 5144 mm 21 4, 5168 mm 31 5, 5171 mm 41 6,, 13142mm 25,, 6127 mm 51 12, 8106 mm Fig. 4 Postlarvae of the tonguef ish 1. 32day2old postlarva, 5144 mm in total length day2old postlarva, 5168 mm in total length day2old postlarva, 5171 mm in total length day2old postlarva, 6127 mm in total length day2old postlarva, 8106 mm in total length ,, mm 2. 24, mm Fig. 5 Juveniles of the tonguef ish day2old juvenile, formation of the dorsal fin and anal fin rays, mm in total length day2old juvenile, eudipleural on both eyes, mm in total length day2old juvenile, initial migration of the right eye, mm in total length.,,, ( 5 : 2 ; 6 : 1) ,, mm, mm

7 1 :, : 125, 96, 4, 8,, ( 7 : 2 ; 6 : 4) , 21 25, 31 27, 41 29, Fig. 6 Front view of the tonguef ish, illustrating migra2 tion of the right eye day2old juvenile, eudipleural in both eyes day2old ju2 venile, initial migration of the right eye day2old juvenile, right eye migrating to the edge of the head day2old juvenile, migration of the right eye completed., 3118 mm,, 4,, 7 ( 5 : 3 ; 6 : 2) , mm, mm, 1138 mm Fig. 7 Juveniles and f ingerlings of the tonguef ish,, 1/ 2, ( ),, 1. 27,, mm 2. 29,, mm 3. 57,, 25192mm 4. 79,, mm day2old juvenile, right eye migrated to the edge of the head, mm in total length day2old juvenile, migration of the right eye completed, mm in total length day2old fin2 gerling, scales appeared, mm in total length day2old fingerling, scales developed completely and reached mm in total length. ( 7 : 1 ; 6 : 3) , mm ( mm, n = 8), , mm ( mm, n = 8) mm ( mm, n = 10),, mm ( mm, n = 10)

8 8 9 50,,, 6 ( 7 : 3) , mm ( mm, n = 10),, mm ( mm, n = 10),,, 3, ( 7 : 312 (1955), 4), ( ),, , Cynoglossus gracilis mm, mm Cynoglossus lighti mm Cynoglossus purpureom aculat us ,, , 1 4 (, 1988a) 311 7, , 100, ( 1),, 1 Table 1 Characters of Cynoglossus spp. in the early development stages Species Water Incubation Egg size Number of temperate period Chorion (mm) oil globules ( ) (h) Number of York crown2like larval fin References S H 1 ( The current study) C. semilaevis HN H 1 Fujita and Uchida, 1957 C. robust us S H 1 C. abbreviat us Fujita and Takita, 1965 ; al., 1986 Fujita et C. arel C. monopus C. joyneri C. puncticep S H 2-3 Ramanathan, et al., S H 2-4 Ramanathan et al., S H 2 Yang et al., 1983 ; Minami, Balakrishnan et al., 1974 C. li da 2 Balakrishnan et al., 1974 C. cynoglossus 2 Balakrishnan et al., 1974 C. bilineat us C. kopsii 2 Ahlstrom et al., Ahlstrom et al., 1984 C. capensis S H 2-4 Ahlstrom et al., 1984 C. macrostom us 2 Ahlstrom et al., 1984 S : HN : H : S : Smooth. HN : Hexagonal Network. H : Homogeneous.

9 1 : 99 2 Table 2 Comparison of the ontogenetic characters of Cynoglossus semilaevis with other Pleuronectiformes Species Newly hatched larva Water temperate ( ) Total length (mm) Standard length (mm) Postlarva Total length Standard length (mm) (mm) ( ) Growth rate Development in size References time (mm/ d) (day) Cynoglossus semilaevis C. robust us C. abbreviat us C. arel ( The current study) Fujita and Uchida, 1957 Fujita and Takita, 1965 ; Fujita et al., 1986 Ramanathan et al., 1979 C. monopus Ramanathan, et al., 1979 Paralichthys olivaceus Zebrias zebra Cleiyhenes herzenstei ni Zhang et al., Zhang et al., Jiang, 1980 Hippoglossoi des robust us Mito, 1963 L i manda punctatissi ma Mito, 1963 L i manda Mito, 1963 herzenstei ni Platichthys stellayus Mito, 1963 Microstom us achne Mito, 1963 Pleuronichthys cornut us Mito, 1963

10 Table 3 Characters of metamorphosis period of Cynoglossus semilaevis and other Pleuronectiformes Species Cynoglossus semilaevis ( ) Right (left) eye began migration Development stage Days after hatching Total length (mm) Metamorphosis finished Days after hatching Total length (mm) ( ) Time of metamorphosis J uvenile period (Day) References ( The cur2 rent study) J uvenile Fujita et al., 1986 C. abbreviat us Zebrias zebra Paralichthys olivaceus J uvenile Zhang et al., 1965 Postlarval Zhang et al., 1965, 40, (1983) , ( ),, ( 60 ),,,, ;,,, 2-3, 313, sp. B rachionus plicatili, ( 5-10 ind1/ ml),,,,, ( , mm) (, 1957), 48 h ( 4132 mm) (, 1995) :, 6 ( 4154 mm) , 115, 8-10 ( mm),, 2, (, 1965, 1986), 3, 13 h ( mm), 5144, 2 ( 5141 mm) mm mm,, 4 ( 5168 mm) mm/ d,,, (24, ( Paralichthyidae Soleidae mm), (5100 mm) Pleuronectidae) Chlorella,, Hi ppoglossoides robust us 4 ( 2) 3,,,,

11 1 :, 315, mm, (317 (,, 1986) ( Zebrias zebra) ( 1955), 1, 1965),, 24,,,,, 29 Paralichthys olivaceus,, (19, 8125 mm),, ; 3, (, 1965), 5 (79 ), 4 (, 1986), (, 1980b ;, 1995) 1965) ( 3), ( References),,,,,,, h, (8-10 ),, 13 h (2 ),,, 4, (12 ), 6,, 1 ( Marine Fisher. Res. (9) : ( In Chinese)., 1965), ( S parus m acrocephal us),2,,,4, (, 1980a), ( Platycephal us indicus) 4, (, 1980b), ( S ardinella z unasi ) (, 1981) ( Konosi rus punctat us) (, 1965) ( Thryssa kam m alensis) (, 1982) ( Engraulis japonicus ) (, 1984 ) ( T richiurus lept urus) (, 1981) ( Scomberomorus niphonius) (, 1966) (, Ahlstrom EH, Amaoka K, Hensley DA, Moser GH, Sumida B Y, Pleuronectiformes : development. In : Moser HG, Richards WJ, Cohen DM, Fahay MP J r, Kendall AW, Richardson SL ed. Ontogeny and Systematics of Fishes. Special Publication Number 1, American Society of Ichthyologists and Herpetologists. Lawrence, KS : Allen Press, Balakrishnan KP, Devi CBL, Larvae of some flat fishes from a Tropical Estuary. In : Blaxter J HS, ed. The Early Life Fistory of Fish. New York : Springer2Verlag Berlin Heidelberg, Blaxter J HS, Hemple G, The influence of eggs size on herring larvae ( Cl upea harengus L. ). J. Cons. Perm. Int. Explor. Mer. 28 : Chen ZR, Zhang XW, A description of the morphological charac2 ters of the eggs, larvae and young of Cl upanodon punctat us ( T. & S. ). Oceanologia et Limnologia Sinica 7 (3) : ( In Chinese). Deng J Y, Meng TX, Ren SM, Qiu XY, Zhu J Y, 1988a. Species com2 position, abundance and distribution of fishes in the Bohai Sea. Deng J Y, Meng TX, Ren SM, 1988b. Food web of fishes in the Bohai Sea. Marine Fishe. Res. (9) : ( In Chinese). Fujita S, Uchida K, Egg development and prelarval stages of a sole, Cynoglossus robust us G nther. The Sci. Bull. Fac. Agr. Kyushu Univ. 16 (2) : ( In Japanense). Fujita S, Takita T, Egg development and prelarval stages of a sole, A reliscus t rigram m us ( G nther). Bull. Japn. Soc. Sci. Fish. 31 (7) : ( In Japanense). Fujita S, Kitajima C, Hayashida G, Induction of ovarian matura2 tion and development of eggs, larvae and juveniles of the tongue2 fish, Cynoglossus abbreviat us, reared in the laboratory. Japn. J. Ichthyol. 33 (3) : ( In Japanense). Jiang YW, The early life history of Flaice, Cleiyhenes herzen2, stei ni nese). 101 (Schmidt). Marine Fisher. Res. (1) : ( In Chi2 Jiang YW, Wan RJ, 1988a. Studies on morphology and developmental characters in early stage of Cynoglossus semilaevis G nther in the Bohai Sea. Marine Fisher. Res. (9) : ( In Chinese).

12 Jiang YW, Wan RJ, 1988b. Reproductive behavior and spawning ecol2 ogy of Cynoglossus semilaevis G nther in the Bohai Sea. Marine Fisher. Res. (9) : ( In Chinese). Jiang YW, Wan RJ, Chen RS, Liu YL, Chen GW, Zhang SB, Fan DD, Fang H, Studies on technique of artificial fry rearing of Cynoglossus semilaevis G nther in the Bohai Sea. Marine Fisher. Res. (14) : ( In Chinese). Li SZ, Wang HM, Fauna Sinica Ostichthyes Pleuronectiformes. Beijing : China Science Press, 68, 334 ( In Chinese). Minami T, The early life history of a tongue fish Cynoglossus joyneri. Bull. Jap. Soc. Sci. Fish. 49 (5) : ( In Japa2 nense). Mito S, Pelagic fish eggs from Japanese waters, Echeneida and Pleuronectida. Japn. J. Ichthyol. Japanense). 11 ( 3/ 6) : ( In Ramanathan N, Natarajan R, Flat fish eggs, larvae and their de2 velopment. A quacult ure 18 : Ruan HC, Studies on the eggs and larvae of Engraulis japonica. Studia Marine Sinica 22 : ( In Chinese). Sha XS, He GF, Zhang XW, A description of the morphologi2 cal characters of the eggs and larvae of the blue spotted mackerel, Scomberomorus niphoni us ( Cuvier & Valenciennes). Oceanol. Limnol. Sin. 8 (1) : 1-8 ( In Chinese). Sha XS, Ruan HC, He GF, A description of the morphological characteristics of the eggs and larvae of the hairtails, Trichi urs haumela ( Forsk l). J. Fish. China 5 (2) : ( In Chi2 nese). Yang DL, Wu GZ, Pang HY, The morphology of the early stage of Longuefishes, Cynoglossus semilaevis G nther and C. joyneri G nther, in the Bohai Bay. Marine Sciences ( 2) : ( In Chinese). Yin MC, Fish ecology. Beijing : China Agriculture Press, ( In Chinese). Zhang XW, He GF, Sha XS, A description of the important morphological characters of the eggs and larvae of two flat fishes, Paralichthys olivaceus ( T. & S. ) and Zebrias zebra (Bloch). O2 ceanol. Limnol. Sin. 7 (2) : ( In Chinese). Zhang XW, He GF, Sha XS, 1980a. Morphological studies of the eggs, larvae and young fish of the black porgy, S parus macro2 cephal us (Basilewsky). Acta Zool. Sin. 26 (4) : ( In Chinese). Zhang XW, Sha XS, He GF, Song LQ, 1980b. A description of the morphological characters of the eggs and larvae of the flathead fish, Platycephalus indicus. Oceanol. Limnol. Sin. 11 (2) : ( In Chinese). Zhang XW, Sha XS, Chen ZR, Ruan HC, He GF, Observations on the eggs and the larvae development of a Clupeid Harengula z u2 nasi Bleeker. In : China Ichthyological Society ed. Transactions of the Chinese Ichthyological Society ( 1 ). Press, ( In Chinese). Beijing : China Ocean Zhang XW, Chen ZR, Ruan HC, He GF, Sha XS, On the de2 velopment of the eggs and larvae of Thrissa kam malensis and Thrissa mystax. Acta Zool. Sin. 28 ( 2) : ( In Chi2 nese). Zheng BS, Cynoglossidae. In : Zhang CL, Cheng Q T, Li SZ, Zheng WL, Wang WB ed. Fishes of the Yellow Sea and Bohai Sea, China. Beijing : Science Press, ( In Chinese).,, (3) : ,,,,, 1988a.. 9 : ,,, 1988b.. 9 : , ( Cleiyhenes herzenstei ni Schmidt). 1 : ,, 1988b.. 9 : ,, 1988a.. 9 : ,,,,,,,, : ,, :, 68, 334., : ,,, (1) : 1-8.,,, (2) : ,,, : , :, ,,, (2) : ,,, 1980a.. 26 (4) : ,,,, 1980b.. 11 (2) : ,,,,, :. ( ). :, ,,,,, (2) : , :,,,,,.. :,

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