Bolm Inst. oceanogr., S Paulo, 40(1/2):55-78, 1992

Distribution and abundance of carangid larvae in the southeastetn . Brazillan Bight during 1975-1981

Mario KATSURAGAWA & Yasunobu MATSUURA

Instituto Oceanográfico da Universidade de São Paulo (Caixa Postal 9075, 01065-970 São Paulo, SP, Brasil)

• Abstract: Distribution and abundance of the carangid larvae collected during ten survey cruises in the southeastem Brazilian Bight were analyzed. TrachuTUS fathami was the most abundant in the surveyed area with 58.54 % of total carangids taken, followed by chrysurus with 15.22 % andDecapterus punctatus with U.17 %. Larvae of T. lathami and D. punctatus were found alI over the while the distribution of C. chrysurus larvae was limited to the coastal region. Most of the species occurred alI year-around but remarkably during spring and summer. This intensive spawning period of most carangid species seems to be related to the seasonal variation of the hydrographic structure of the southeastem of Brazil

• Descriptors: , Ichthyoplankton surveys, larvae, Population number, Seasonal distribution, Abiotic factors, RN ''Prof. W. Besnard", Southeastem Brazilian coast.

• Descritores: Carangidae, Levantamentos ictioplanctônicos, Larvas de peixe, Número da população, Distribuição sazonal, Fatores abi6ticos, N/Oc. ''Prof. W. Besnard", Brasil: costa sudeste.

Trachinotus. Global nominal catches of the carangids in Introduction 1987 was abollt 240.93 thousand tons (FAO, 1989). They are also important for sport fishing (Seriola spp.) in Carangid inhabit marine and estuarine waters in California (Sumida et ai., 1985) or for fishculture (Seriola tropical, subtropical and temperate regions of the world quinqueradiata) in Japan (Fukuhara, 1987). ln the marine (Bannikov, 1987). They are active swimmers and live ecosystem they are important prey species of many in school or smalI groups (Berry & Smith-Vaniz, 1978). predators such as tunas and tuna-like species (Bailey, 1989; The group is heterogeneous, including genera of quite Shaw & Drullinger, 1990). different shapes and appearances, from elongated and ln Brazilian waters the study on distribution and fusiform to high and lateralIy compressed body (Ginsburg, abundance of carangids is scarce. Menezes & Figueiredo 1952). The carangids are constituted by approximately 32 (1980) elaborated an identification manual of marine fishes genera and 140 species (Smith-Vaniz, 1984; Gushiken, which includes information of occurrences and 1988), from which 15 genera occur in the southeastem distribution of 30 carangid species. Saccardo (1980; 1987) Brazilian waters (Menezes & Figueiredo,1980) : investigated the morphology, reproduction, distribution, Trachinotus, O/igop/ites, Parona, Naucrates, Serio/a, and the abundance of Trachurus lathami in southem A/ectis, Caranx, Ch/oroscombrus, Decapterus, , Brazil. Zavala-Camin (1981; 1986) showed occurrences of Pseudocaranx, , , Trachurus, and Uraspis. carangids species in stomach contents of tunas and related Many of them are commercialIy exploited, specialIy species in southem Brazilian waters. Cunha (1987) those of the genera Trachurus, Caranx, Decapterus and descnbed the occurrence, distribution, and bioecology of Trachinotus species in intertidal zones from Rio Grande do Sul to Espírito Santo. Scorvo-Filho et ai. (1987) Contr. no. 732 do Inst. oceanogr. da Usp. described an occurrence of juveniles Trachinotus species 56 Bolm Inst. oceanogr., S Paulo, 40(1/2), 1992 in the Ubatuba region. Very üttle is known about egg and Sampling methods larval abundance and distribution patterns of carangids in Brazil with exception the distribution and abundance of Plankton samples were collected by Bongo nets with Chloroscombrus chrysurus larvae ai ong the southern digital flowmeters mounted at the center of mouth (Posgay Brazilian coast described by Weiss et ai. (1976). el ai., 1968). The samplingmethod used was that Knowledge obtained from ichthyoplankton survey descnbed by Smith & Richardson (1977). Two nets with can provide important information about the different mesh sizes (0.333 mm and 0.505 mm) were used, potential of a species (Richards, 1985). Presence of a large but only the plankton samples collected with the 0.505 mm number of eggs and larvae in a certain area indicate the net were considered in this study. Double oblique hauls pre~ence of adult schools. Thus, the study of distribution were made and sampling depth varied according to station and abundance of eggs and larvae can provide estimate on depth, but maximum for 200 m depth. ln stations the fluctuation of adult stocks as well as changes in shallower than than 60 m the net was lowered twice elose spawning area and season due to changes in oceanographic to the bottom to increase the volume of water mtered. 'lhe conditions (Ciechomski, 1981). ship speed was maintained in 1.5 to 2.0 knots to keep the ln this paper we present informations on occurrence wire angle in about 45°. 'lhe samples of carangid larval groups, the seasonal distribution collected with the nets were preserved in a buffered patterns of the most abundant species, and the influence of formalin 10 % solution. abiotic factors on them in the southeastem Brazilian At each sampling station a routine hydrographic Bight. observations were made using the Nansen bottle with reversing thermometers, !lDd temperature and salinity of discrete depth were measured Temperature and salinity at 10 m depth of each station were considered as Material and methods representative of the upper mixing layer, where most of fish eggs and larvae were found. Surveyed area

'lhe samples of carangid larvae were collected Laboratory procedures during ten survey cruises carried out in the southeastem Brazilian Bight (SEBB) with the R/V "Prof. W. Besnard" of the Instituto Oceanográfico da Universidade de São ln the laboratory, the zooplankton volume was Paulo and with the R/V "Cruz deI Sur" of measured by displacement method (Kramer et ai., 1972). PDP-SUDEPE (Table 1). Thesurveyarea extended from All flSh eggs and larvae were frrstly sorted' out from Cabo Frio (23 OS) to Cabo Santa Marta Grande (29 OS), plankton samples using Bogorov plate, and then the larvae covering entire continental shelf up to 200 m isobath (Fig. were elassified into group. 'lhe carangid larvae were 1). ln the cruises 1,2, 3, 4,5 and 7, we extended two identified into species leveI according to the morphological observation lines for 260 km from the coast in Cabo Frio characters of each species (Katsuragawa, 1990). and in Cabo Santa Marta Grande to cover the main axis of Larval abundance was estimated using the following the Brazil Current along the shelf break. ln the cruises 10 equation: and 12, the surveyarea was limited up to the 100 m isobath line on the continental shelf. y= D.X V

Table 1. Summary of the oéeanographlc crulses conducted between 1975 and 1981 ln the southeastern Brazillan Blght

No. OF CRU I SES DATES VESSEL STATlONS SAMPLING AREA

FINEP 1 NOV-DEC/1975 R/V "Prof. W. Besnard" 140 Cabo Frio (RJ) - Cabo de Santa Marta Grande (SC) FINEP 2 JAN/1976 RlV "Prof. W. Besnard" 140 Cabo Frio (RJ) - Cabo de Santa Marta Grande (SC) FINEP 3 MAY/1976 R/V "Prof. W. Besnard" 140 Cabo Frio (RJ) - Cabo de Santa Marta Grande (SC) FINEP 4 SEP-OCT/1976 R/V "Prof. W. Besnard" 140 Cabo Frio (RJ) - Cabo de Santa Marta Grande (SC) FINEP 5 DEC/1976 R/V "Prof. W. Besnard" 140 Cabo Frio (RJ) - Cabo de Santa Marta Grande (SC) FINEP 6 JAN-FEB/1977 R/V "Prof. W. Besnard" 112 Cabo Frio (RJ) - S. Francisco do SUL (SC) FINEP 7 JAN/1978 R/V "Prof. W. Besnard" 134 Cabo Frio (RJ) - Cabo de Santa Marta Grande (SC) FINEP 10 JAN-FEB/1979 R/V "Cruz de L SUr" 90 Cabo de S. Tomé (RJ) - Cananéia (SP) FINEP 11 JAN/1980 R/V "Prof. W. Besnard" 42 Cabo Frio (RJ) - FLorianópoLis (SC) F1NEP 12 JAN/1981 R/V "Prof. W. Besnard" 46 Cabo Frio (RJ) - FLorianópoLis (SC) KATSURAGAWA & MATSUURA: Carangid larvae: Distribution and abundance: Brazilian Bight 57

wbere Y = number of larvae per square meter of sea Decapterus punctatus were predominant, representing 83.9 surface, D = maximum sampling depth (m), X = number %. Eleven percent of larvae and juveniles were not of fisb larvae taken, and V = volume of water filtered (m3) . identified to any taxon. But, V = a.n.c, wbere a = area of mouth opening (0.2827 m2), n = number of rotatioDS of flowmeter and c = calibration factor of flowmeter. Rougb scad, Trachurus lathami

Larvae and juveoiles of rougb scad, r. lalhami, were the most abundant (58.5 %) among all carangids studied Results in this reporto The distribution of this species extended from the coast (16 m depth) to the sbelfbreak, covering the Table 2 shows tbe number of carangid larvae and entire continental sbelf (Fig. 2). However, we can notice juveniles collected during ten survey cruises in tbe that the higbest density of larvae (more than 20 larvae m2) soutbeastem Brazilian Bigbt. Five taxon were identified at was localized in tbe neritic region. A relation between the species leveI (Trachurus lathami, Chloroscombrus occurrence and station deptb sbowed that 64.5 % of chrysurus, Decapterus punctatus, and positive statioDS and 85.5 % ofr. lathami larvae were found Selene vomer) and four at leveI (Oligoplites, Caranx, in the 4O-100m depth zone (Fig.4a). Occurrenceoflarvae Seriola and Trachinotus). From a total of 12,600 carangids, and juveoiles in waters off the continental sbelf was rare. A Trachurus lathami, Chloroscombrus chrysurus and similar tendency was observed for other carangid larvae.

FINEP BASIC STATION PLAN

o o o o o o o o o o o o o o o

o Stations

o o o o o o

48° 400

Fig. 1. FINEP basic station plan. ranging frem Cabo Frio (23°S) to Cabo de Santa Marta Grande (2go$) in the .southeastern BrazUian Bight. ~

Table 2. Total number of carangld larvae anallzed per crulze

CRU I SES Trachurus Chloroscombrus Decapterus Selene Selme Oligoplites • Caranx Seriola Trachinotus . Others TOTAL lathami c1uysurus punclatus setapinnis vomer spp spp spp spp carangids

FINEP 1 658 W 202 2 o o o o 459 1446 FlNEP 2 301 114 115 47 18 23 5 o 153 m FINEP 3 5 W 11 o o o o o o 21 134 FlNEP 4 3942 7 o o o o 2 o 123 4075 FINEP 5 785 365 856 19 3 o 2 o o 206 2236 FINEP 6 107 195 139 22 2 4 12 o o 145 626 FINEP 7 1037 888 132 67 3 27 12 o o 199 2365 ~ FINEP 10 133 2 9 2 2 O 4 o O 1 153 o FlNEP 11 113 76 32 3 4 38 O 2 O 20 288 ã FINEP 12 268 83 30 3 3 22 16 o O 75 500 ri o R TOTAL 7376 1918 1533 165 35 114 51 5 1402 12600 ~ X TOTAL 58.54 15.22 12.17 1.31 0.28 0.90 0.40 0.04 0.01 11.13 ~ CI'l "I:j c»e. wo -~ ~.... ~ KATSURAGAWA & MATSUURA: Carangid larvae: Distribution and abundance: Brazilian Bight 59

ln November-December 1975 two large which 92.3 % oflarvae were collected Only 1.4 % oflarvae concentrations of larvae were found: one near were found in stations with more than 50 m. Florian6polis and another in São Sebastião-Ubatuba Distribution of C. chrysurus larvae extended from region (Fig.2a). ln January 1976 only one large Cabo Frio to Florian6polis with large variation in concentration was found along llha Grande coast. ln May distribution patterns among different cruises (Fig. 5). C. 1976 no large aggregation was found, and the distribution ChrySUlUS larvae were found mainly in the coastal stations was restricted to small number of larvae in the region between São Sebastião and São Francisco do Sul in all between Santos and São Francisco do Sul (Fig.2c). late-spring and summer cruises. Exceptions to this pattem ln the early spring cruise (Sep-Oct. 1976) four large occurred in both January 1976, when the occurrence concentrations of larvae werefound along the coast from extended further north, and Jan-Feb. 1979, when no São Sebastião to Cabo Santa Marta Grande and the significant occurrence was found. ln autumn (May 1976) abundance of T. lathami larvae was the largest of the ten the larval distribution was limited to the coastal region cruises (Fig. 2d). ln the following cruise (Dec. 1976) total between Cabo Frio and Paranaguá. ln September­ abundance decreased and two areas of larval October 1976 on1y one positive station was found dose to concentration were found: one in northeast of Rio de Janeiro, indicating that no significant spawning Florian6polis and another along Ilha Grande coast (Fig. activities were going on during late spring and summer that 2e). During the cruise of January-February 1977 no large year. concentration was recorded. On1y small numbers of larvae Temperature variation in positive stations of C. were found over the continental shelf. ln January 1978 a chrysurus was between 16.1 and 28.7 °c with mean large concentration of larvae was recorded along the coast temperature of24.8 °c (s = 2.10). Most of positive stations of Ilha Grande, but no significant number of larvae was had temperature higher than 22°C, 73.9 % of them were found in southem part of the surveyed area (Fig. 2g). found between 24° and 27°C, in which 90.0 % oflarvae were For the last three cruises conducted in taken (Fig. 3b). This result shows a slightly higher January-February 1979, January 1980, and January 1981, temperature range of this species in relation to T. lathami. only the area inside the 100 m isobath line has been investigated. ln January-February 1979 (Fig. 2h) no large Salinity variation was between 32.75 and 36.89 psu concentration was observed but larvae were present in all with mean salinity of 34.70 psu (s = 0.92). Most positive surveyed area. ln January 1980 the number of positive stations stayed between 33.50 and 36.00 psu, in which 86.5 stations decreased further and a large amount of larvae % oflarvae were taken (Fig. 3b). were found along the coast of Rio de Janeiro State (Fig. 2i). ln 1981 the distribution pattem of larvae was similar to that observed in 1979 but a large concentration of larvae was found in front of Rio de Janeiro in offshore waters Round scad, DecaptelUs punctatus (Fig.2j). T. lathami larvae were found in a wide temperature range between 11.5 °c and 27.5 °c (Fig. 3a), the mean DecaptelUs punctatus was also abundant and temperature at 10 m depth of all positive stations was 22.2 accounted for U.2 % of total carangids col1ected (Table °c (5 = 2.67). The majority of positive stations (85.8%) and 2). This species showed a wide distribution over the of the total catch (94%) were observed within the continental shelf, similar to that observed for T. lathami, temperature range of 18-25 oCo The salinity at 10 m depth but induding some positive stations out of shelf break off of all positive stations varied from 32.86 to 37.67 psu but Cabo Frio. Occurrence of positive stations was 88.2 % of positive stations and 95.9 % of larvae were found concentrated in depth zone between 20 m and 60 m, in in salinity range of 33.5-36.0 psu. which 94.4 % oflarvae were taken (Fig. 4c). The occurrence of DecaptelUS punctatus larvae showed a large variation in terms of abundance and horizontal distribution (Fig. 5). They were more abundant during austral summer (from December to , ChloroscomblUs ChrySUlUS February) cruises and less abundant in autumn and early spring cruises. Their horizontal distribution patterns during late-spring and summer cruises were similar to The atlantic bumper, ChloroscomblUs ChrysulUS, was those observed for T. lathami, but more sporadic over the the second most abundant species representing 15.5 % of continental shelf. D. punctatus larvae were always all carangids collected (Table 2). Larvae of this species abundant in coastal areas off Paranaguá during eight were found in all cruises but their distribution was limited summer cruises. They were also found in coastal areas off exclusively to coastal stations (Fig. 5). They were collected Santos and São Sebastião. The largest concentration of D. at depth range from 15 to 75 m (Fig. 4b) and mean depth punctatus larvae was observed in coastal areas off of all positivestations was 31.5 m (s = 14.9). Most of Paranaguá between Santos" and São Francisco do Sul in positive stations were found in the 15-30 m depth zone, in December 1976. 60 Bolm Inst. oceanogr., S Paulo, 40(1/2),1992

FIg. 2. Olstrltulon and abundance (no. of Iarvae/m2) of Trochurus lathami 18IV8e. a) November and Oecember 1975; b) January 1976. KATSURAGAWA & MATSUURA: Carangid larvae: Distribution and abundance: Brazilian Bight 61

c 200m

o o o ~ _ -

0/ ' o ..~_ .. o o , , o ,, o , • o I , , o o o , o o 0 ... _ ___ _ " , o o o o ,6 o o , O O O , /\AV 1976 , " TJLacJ,,,ULb O , ~ . o,'" O STAT/ONS O ,, b no. lo,voe/ m' o 0,1 5,0 o ~ - o 5,1 -10,0 o o ,: • , o o , 10,1 -20,0 o • 20,1 -30,0 o , o o , o • > 30,0 o •

500 W 48° 46° 44° 42°

d 200m

,, o • o • o 0 ... -0 , • o , \, • • • • " o • O ... f • Q.. _ .... o o ... ' .. • • ,, • •• , o • o • • ••• • , o • • ,, " • o " , • ••• 0,..'" _ __ _ " • • o •• •• ,6 o , SEP ,-OCT. 1976 • , • ," • , , • . ,' o STAT/ONS ,, • • ,b no. lo,voe / m' • • • 0,1 - 5, 0 • • 5,1 - 10,0 • .,o" • 10, 1 - 20,0 . ,/ • 20,1 - 3 0,0 .. ' • > 3 0,0

Fig. 2_ Distribution and abundance (no. of larvae/m2) of TrachulUs lolhami larvae. c) May 1976; d) September and October 1976. 62 Bolm. Inst. oceanogr., S Paulo, 40(112),1992

e

• • o • •

o • 0 #' .... 0 o o o "o- -,, '" o • , o o • o , o o . : o o o o o o o o o " o • o o ' o o o o o 0 ...... - __ _ " • o o o ,6 • o ,; • o , o DECE"IER 1976 • o , o ," r""ohuIuu Whawu: • o o,, ' o STATlONS o , • ,b no. lorvoe/m l • o 0,1 - 5,0 o , o , 5,1 -10,0 o , • , o , 10,1 -20,0 . • 20,1 -30,0 o , o , o o • o • > 30,0 500 W 48° 460 44° 42°

f

o o o o o o • o o o o o • o o o o o o o o O ,;;" ,r ' JAN.-FEB. 1977

t r""chu .... ~ o ) 27° I'"'' o o STATlONS (; no. lorvoe / m Z • 0,1 - 5,0 ,, 5,1 - 10,0 ,, • , 10,1· - 20,0 , .20,1• - 30,0 > 30,0 • 29° 480 44° 42° 40°

Fig. 2. Dlstrlbutlon and abundance (no. of larvae/m2) of TrachunlS latham; larvae. e) December 1976; f) January and February 19n. KATSURAGAWA & MATSUURA: Carangid larvae: Distribution and abundance: Brazilian Bight 63

9 200m

, o o ,

o 0 ... - 0 , o '" \ .... ' o o

o • o o • • • o , o • , o o • • , •o • o • ," o o , o o ,­ o .. o , o 0, ... ____ ' o • o • o ,6 o , o ,o o o JANUARY 1978 o o ,,.' o , , Tllo.dw.• "" l4tham< • o 27° o, ' o , o STAT/ONS o , • ", no. lorva" / m I 0,/ - 5,0 o ~, • o , 5,/ -/0,0 o o , • o ,, o /0,/ -20,0 o o o • 20,/ -30,0 o , o , o o • , o > 30,0 o • 29° 440 420 400

o, ,' o o 9" ...... o ,- - õ "

p'

240

o , o o ., JAN . -FEB . 1979 • • ' 0 ',",' o ' T""o h"""" t a.tho.mi. o '- , • • o, , o STAT/ONS o o no. larva" / mW o • .... ~ , ~ ,Q...... o 0,/ - 5,0 o ," - .. ;I • • , o • 5,/ - /0,0 9' 10,1 ~20,0 .20,/• - 30,0 260 • > 30,0

Fig. 2. Distribution and abundance (no. of larvae/m2) of Trachurus lathami larvae. g) January 1978; h) Jamllry and February 1979. 64 Bolm Inst. oc:eanogr., S Paulo, 40(1/2), 1992

,, 1 23°5 ,~ I • • • I , . . -, , • • ,- I • , . ' _., - o , o • ,, o o , o , 25° , • ,, o , , , : • , , o , o ,, I JAIIUARY 1980 o ,, ,, T/14dw.w.6 ~ . , 27° o , I o , o STATIONS no. larva. I ",. o , • 0,1 - 5,0 ,, • 5,1 -10,0 , 10,1 -20,0 , • -30,0 I • 20,1 > 30,0 • 29° 500 W 48° 46° 44° 42° 400

J

, ---_.. \ ...... • o o

o • .' • o /,"-._' ,'-" -'

o ,,;// o '

JAIIUARY 1981 ,~ o o, o STAT/ONS no. lorvoe/m' • 0,1 - 5,0 • 5,1 - 10,0 • 10,1 - 20,0 IÓOm .20,1 - 30,0 • > 30,0

Fig. 2. Dlstributlon and abundance (no. of larvae/J) 'of 1'rriChurus latham; larvae. I) January 1980; j) January 1981. § a) T. lafhami ~ »~ i 1 I 1 P:o I I I ~ ~ ~ I I [ I i. Q. ~ ~ t::) ln· ~ I I ct. L J ~ [

I I I I 1/ I /0 /4 /8 22 26 30 32 34 36 38 J t:I:I TEMPERATURE (Oe) SAL/N/TY (psu) t t:I:I Fig. 3. Temperature and sallnlty ranges obtalnecl from ali positive statlons for three specles: a) Trachums ti... latham~ b) Chloroscombrus chrysurus, and c) Decapterus punctatus.

~ 66 Bolm Inst. occanogr., S Paulo, 40(1/2), 1992

80 80 .J Trachuru. I.Ihaml p o 80 8 80 I .. T 40 I o V C E C 30 40 U 8 A T A A I! T 20 N I C o 20 E N 8 10

o O 10 30 60 70 90 110 130 160 170 190 OEPTH (m)

80 80 bJ Chloroacombru. chryaurua p o 60 8 60 I .. T 40 o I C V C E U 30 40 A 8 A T E A N T 20 C oI 20 E N 8 10

O O 10 30 60 70 90 110 130 160 170 190 OEPTH (m)

60 80 c) Dtlcapttlrua punctatua p o 60 8 60 I .. T 40 o I C V C E U 30 40 A 8 A T E A N T 20 C oI 20 E N 8 10

O O 10 30 60 70 90 110 130 160 170 190 OEPTH (m)

_ POSo STATIONS _ ~ OCCURRENCE

Fig. 4. Proportlon of number of larvae by depth zone and number of positive stations of three species: a)Trachwus lathami, b) Chloroscombrus chrysurus, and c) DecapteTUs punctatus. Ali larvae taken at statlons deeper than 200 m are presented at the last column. KATSURAGAWA & MATSUURA: Carangid larvae: Distribution and abundance: Brazilian Bight 67

a 200m

00 o o o o o o - o o 00 ~ - , o o --- .. .,'0 06 o 06 o o o o , o. o o , o o o o o 06 o o o " o o o o o o o o o " 06 o 06 o o o o , o o... ~ .. _ ...... ' NOV. -OH . 1975 •• o o o • o o , • Ch.lo 30,0

b 200m , , o o o o o 06 o o o o .. _ .o •• o o , \,,_ o 06 o o o .. ' o o .~_ .. " o 00 o ' o , 00 tO 06 , o o o / ' o o o o o o o o 06 o , " o o , o , o o o o o , 06 , JANUARV 1976 0, ... ':" .. _ ...... o • Cht"'Lo&c.OrtTb~6 Chl!.y6U.W~ o 06 o • VecapÚw..l pUl1ct4.tu4 06 o • Sclera' .UttlpullI.(.6 • Se tC.lH~ vcmc. 'I.

O,' o O STAT/ONS o, no. larvae / m' O t • ~I - ~O O o • • • • 5,1 - 10,0 o o o ," o • ••• 10,1 - 20,0 o , • • Â • 20,1 - 30,0

o • • Â • > 30,0

Fig. 5. Dlstrlbutlon and abundance (no. of Iarvae/m2) of Chloroscombrus chrysurus, Decapterus punctIItUS, Selene setllpinnis and Selene vomer larvae. a) November and December 1975; b) January 1976. 68 Bolm Inst. oceanogr., S Paulo, 40(1/2), 1992

c 200m , ,

o o o I o 06 o o I o 06 o 06.. __ O, .. - (I '" o "" o o o o o o O,, o ,Q...... o o o o , o • o ,, o o , o o I o o o o o o o • o o o o o o " o o , o o 0, ... ___ .. " o o /lAy 1976

o o • chto~o.c ...b""" cJvtY'1JJIJU o • Oecapa""" pwtc.útu.o o , o , o o I " o , o,' o , o STAT/ONS o , o ,b no. lorvoe / m t

~ 6 0,1 - 5,0 ... 5,1 -10,0 o o , • o , o 10,1 -20,0 o , o • • , o o 20,1 -30,0 o • • o ... • > 30,0 29°

500 W 48° 46° 44° 42° 40°

d 200m

o o o o o O ... - (I , o o o .. __ ... .. \ ...... '0 o o o o o , o o o , o o o , o o o o , o o , o o , o 06 ,

"

SEP . -QCT. 1976

o o • Ch i M o .comb~ ... CMy...... • Oe capa""" pUl1 C.ta.tlU

o I

O,' o STATIONS ,b no. lorvoe / m'

O 0,1 - 5,0 ... • 5,1 - 10,0 , o o , •• 10,1 - 20,0

O •• 20,1 - 30,0 .... > 30,0

Fig. 5. Dlstrlbutlon anel abunelance (no. of larvae/m2) of Chloroscombrus chrysurus and Decapterus punctatus Iarvae. c) May 1976; d) September anel October 1976. KATSURAGAWA & MATSUURA: Carangid larvae: DistributiOD and abundance: Brazilian Bight 69

e 200m

o .0 o ~ ...... 0\ o .... ' 0 OlA .0 o .0 o, , , ' o Q..., o o to o ~ , " 'IA. • o o o o , I o q6 ao , o 06 o o .. o o DECE"BER 1976 0 ...... ___ " o C hto~o .comb""" CM~''''''' ,6 • Â Dc cap.te ..... pw!c.(4~ 06 .; 06 Se te... ".(ap.u..u.. o .. • Se.tent vomelt o , • o, ' o o , STATlONS o , ., no. larva. / m t o , o t O,, - 5,0 o ~ .. o , I ... 5,1 -10,0 o o , • o • • o , Â 10,1 -20,0 o , o o • 20, 1 -30,0 o • • ~ o , o o o I • • 30,0 o • • .. • >

f

o o o o o OA o .0 o o eo o ao o o o ..... o o o o - o o o o o , o o ao o o , o o o , .' JAN . 'FEB . 1977 • o o o o • C IICo't( ' ~ c"",b"UW d l Jt.y ~ u 'l u,~ o o , o 'f  Vc captc",o PU'1C c"tJ.<. • ~ , • Sefene ~ c tap.uw,(,!I .0 ~, o o o , o o STATIONS ( o no. larva. / mZ , • • O, I - 5,0 • ... • 5 , 1 - 10,0 • .... 10,1 - 20,0 • ~ • 20,1 - 30,0 • ... > 30,0

Fig. 5. Distributlon and abunelance (no. of larvaeJrif) of Ch/oroscombrus chrysurus, DeaIpteTUS punctatus, Selene setapinnis anel Selene vomer larvae. e) December 1976; f) January and February 19n. Bolm Inst. oceanogr., S Paulo, 4()(1J2), 1992 70

9 ~m I

o o 0 ... -0 , 00 o 0... , __ ,' ' .... / 06 o .o o., 0 .... ' 0 Q.. -,' o o o o .c)6 o," o 00 , o o o o , " o o o o , o o , " o o JANUARY 1978 0 ... .. ____ .... o o • ChtoJto.

o , • ~lLn~ ..t.ap..:n.u. o ,. • v",""Jt o I ~lLn~ o , , o o,' o STATiONS o I o I ,b no. lorvo,,/m' • 0,1 - S,O o ~, • • • • S,I - 10,0 o ' • • Â • 10,1 -20,0 o ," o o I o • • Â .20,1 -30,0 o , o o • • Â • > 30,0 o • 500 W 48° 46° 44° 400

h

o p' ,"'-0, / JAN . -FEB. 1979 o o o f ,0 CMMo.eomb·,uo d",y• ..w.. 24° 0 " .-0 • o  ~ 30,0 100 m

Fig. 5. Dlstributlon and abundance (no. of larvae/m2) of Ch/oroscombrus cJuyswus, Decapterus punctatus, Selene setapinnis and Selene vomer larvae. g) January 1978; h) January and February 1979. KATSURAGAWA & MATSUURA: Carangid larvae: Distribution and abundance: Brazilian Bight 71

, I , -' o o I

o , I o ,

, , " .,. , I .,. o JA"UARV 1980 , , • ChtOl10600mblUJ1> olvty6U1U.L6 ... fkoapt2.1UJ1> punc..tl:lW.6 .. , • Set."" 6 • .t.lp.in.u. • Sete. ne vomeJt , ,I o , , I o STATIONS I I , no. larvatl / m' o , • 0,1 - 5 , 0 • • • 5,1 - 10,0 • .... 10,1 - 20,0 , • .... 20,1 -30,0 I • •• > 30,0

480

• J

Fig. 5. Dlstrlbutlon and abundance (no. of IarvaeJm2) of Chloroscombrus chrysurus, DecapteTUs punctatus, Selene seúZpÍII1IiS anel Selene vomer larvae. i) January 1980; j) January 1981. n Bolm Inst. oceanogr., S Paulo, 40(112), 1992

During the May 1985 cruise only small numbers of D. collected in the southem Gulf of Mexico the three most punctatus larvae were found in the region between Cabo abundant species were C. chrySUTUS (54.2 %),D. punctatus Frio and Santos, and during the Sep-Oct. 1976 cruise small (15.8 %) and T. lathami (11.8 %). ln the eastern Gulf of numbers of larvae were found in neritic regions off Mexico three dominanl- species were D. punctatus, C. Paranaguá. chrysurus and T.IOIhami in this (Leak, 1981). We can Temperature variation at 10 m depth of positive observe that the most abundant species changes from area stations of D. punctatus was between 14.7 and 28.4 °C with to area, but larvae of those three species always mean temperature of 24.4 °C (s = 2.29) (Fig. 3c). This predominated in the westem Atlantic corresponding to temperature range was similar to that observed for C. more than 80 % of carangid larvae. chrysurus, but 2.0 °C bigher than that for T. 1000ami. Inside Studying the occurrence of carangid larvae in the the temperature range of Zl-Z7 °C, 87.0% of positive coast of Louisiana State, Shaw & Drullinger (1990) stations and 95.2 % of larvae were found. Salinity variation considered that the blue runner, Caranx crysos, also can be was between 32.96 and 37.00 psu with mean salinity of35.26 included in the dominant group because it was the second psu (s = 0.87). most abundant species after D. punctatus. ln the present study the larvae of C. crysos could not be identified from other Caranx larvae, but the overall abundance of Caranx larvae collected in this area was relatively small (0.4 % of

Atlantlc moonftsbt Selene setapinnis and lookdOWllt all carangids). Thus, we can conclude that they are really Se/ene vomer less abundant in the surveyed arca. This result surprised us, since the adults of C. crysos are common in the area and The larvae of Atlantic moonfish, Se/ene setapinnis considerable commercial catches are known. was less abundant than those discussed above, Low abundances of other genera and species of representing only 1.31 % of total carangid larvae collected carangids in our samples coincided with the results (Table 2). They were collected in alI cruises of obtained in the eastem coast of the North America by other late:-spring and summer and none were found in May and authors (Aprieto, 1973; Leak, 1981; Flores-Coto & Sep-Oct. Their distribution was basically neritic, but in Sanchez-Ramirez, 1989). Two areas of investigations, some cases they were also found in coastal areas and off southeastern Brazilian Bight and Gulf of Mexico, continental shelf (707 m depth). There was no specifically presented a similar assemblage of species composition of important area of occurrence, but they were found the family Carangidae, indicating a zoogeographical sporadically from Cabo Frio to Cabo Santa Marta Grande linkage between both areas. Leak (1981) considered that (Fig. 5). some species were rare in bis study because theymay spawn The larvae of , Selene vomer was less out of the continental shelf or very close to the coast (e.g. abundant than S. setapinnis, representing only 0.28 % of Oligoplites saurus), which could not be sampled by the total carangidlarvae, or 12.5 % of genusSelene. They were sampling system applied. ln the case of Oligoplites this also found only in summer cruises and their distribution assumption is true in the SEBB since they are frequently was more limited to coastal regions, not exceeding the 100 collected in shore line or estuarine region. misobath. Temperature range ofS. setapinnis was from 17.1 to 26.6 °C with mean temperature of 24.4 °C (s = 1.62) and that of S. vomer was from 16.7 to 27.9 °C with mean Spatlal and temporal distributlon of larvae temperature of 23.6 °C (s = 3.47). Salinity range of S. setapinnis was from 34.31 to 37.01 psu with mean salinity of 36.01 psu (s = 0.66) and that of S. vom(!r was from 33.03 to Among 13 known species of the genus Trachurus in 36.50 psu with mean of 35.07 psu (s = 0.87). the world, T. 1000ami is the only one occurring in the westem Atlantic fram the Gulf of Maine to the narthem Argentine (Berry & Cohen, 1974). ln the sautheastem Brazilian Bight they are typical neritic species, occurring Discussion fram depth zone between 50-100 m (Saccardo, 1987). Menezes & Figueiredo (1980) observed that this species did not approach to coastal regioos less than 40 m depth. Relatlve abundance On the other hand the distributions af larvae and juveniles demanstrated that théy could occur from the coast (16 m depth) to as far as off continental shelf. However, the Three carangid species predominated in the main areas of occurrence (85.5 % of positive statians) were surveyed area: Trachurus lathami (58.5 %), found in the same depth zone of adult (40-100 m). Similar Chloroscombrus chrysurus (15.2 %) ánd Decaptems results were obtained in the eastem Gulf af Mexico where punctatus (12.1 %). They are also abundant in other areas larvae were found in depth zone 20-219 m, but major of the western Atlantic. Flores-Coto & Sanchez-Ramirez accurrences were in neritic region (more than 50 m depth) (1989) verified that among eight species of carangid larvae (Leak, 1981). On the Louisiana shelf T. Ialhami larvae KATSURAGA WA & MATSUURA: Carangid larvae: Distribution and abundance: Brazilian Bight 73

were collected in depth zone 10-100 m, but 99 % of larvae period (Conand & Franqueville, 1973) and in the GuIf of were found in waters more than 30 m depth (Ditty & Mexico they were found in June-August period (Shaw & Truesdale,1984). Drullinger, 1990). Presence of Iarvae can be considered as an index of Species of genus Deeapterus occur in neritic and recent spawning in lhe area (Farris, 1961; Aprieto, 1973), oceanic waters in tropica~ subtropical and temperate specially for the pretlexion Iarvae. Thus, we can estimate regions (Stanton Hales, Jr., 1987). ln the western Atlantic the spawning areas and its intensity based on the three species are known: D. maearellus (Cuvier), D. distribution pattem of larvae and their size frequency punetatus (Cuvier) and D. tabl Berry (Berry & Smith­ distribution. From the results obtained in this studywe can Vaniz, 1978), from which D. pune tatus is the most conclude that the spawning of T.lathami in the SEBB takes common in the southwest Atlantic. D. tabl may also occur pIace in all surveyed area, but with two areas of preference, in the southeastem Brazil (Menezes & Figueiredo, 1980), one between Rio de Janeiro and Ubatuba and another but we found onIy larvae of D. punetatus. Larval between Paranaguá and Florian6polis. distnbution pattem of this species was similar to that Spawning season of T.lathami extends all year-round observed with T. lathami from the coast to off continental in the area since the pretlexion larvae were collected from shelf area. This pattem is also observed in other areas all survey cruises. However, it was more intensive in early (Hildebrand & Cable, 1930; Aprieto, 1973; Leak, 1981; spring and summer cruises. Peak abundance was observed Naughton et al., 1986). in the Sep-Oct. auise and the lowest abundance was ln the Gulf of Mexico and the South Atlantic Bight observed in May cruise (autumn). Based on D. punetatus larvae occurred all year-round with seasonal macroscopical observation of adult gonads, Saccardo varlation in different geographical areas (Aprieto, 1973; (1980) observed more limited spawning season from spring Naughton et ai., 1986). For exampIe, the peak spawning to summer with peak spawning in November in the same occurs during spring in the South Atlantic Bight (Fabay, area. On the Uruguayan shelf and northem Argentine a 1975; Johnson, 1978) and during spring-summer in the Gulf short spawning season was observed from November to of Mexico (Leak, 1981). ln the surveyed areathey were March with intensive spawning in December and January found all year- round peaking in austral summer. (Cousseau, 1967; Ciechomski & Weiss, 1973; Ciechomski Distribution of S. setapinnis extends from Nova & Cassia, 1980; Pacheco Tack, 1988). ln the Gulf of Scotia, Canada, to Mar dei Plata, Argentine. Adults of this Mexico T. lathami spawns ali year-round with peak species form schools and live close to the bottom in coastal spawning in Iate autumn and earIy spring (Leak, 1981; region (Berry & Smith-V aniz, 1978). ln the SEBB Flores-Coto & Sanchez- Ramirez, 1989). occurrences ofS. setapinnis larvae were scattered over the Geographical distribution of Chloroseombrus continental shelf which was similar to the results obtained ehrysurus extends from Massachusetts to Uruguay in the in the southern Gulf of Mexico (Flores-Coto & westem Atlantic (Berry & Smith-Vaniz, 1978) and from Sanchez-Ramirez, 1989). They were mainly found during MoiToco to Senegal in the eastem Atlantic (Aboussouan, austral summer in the SEBB, but in low density. Aprieto 1968; Conand & Franqueville, 1973). Habitat of adult (1973) found small number of S. setapinnis larvae during extends from coastal waters to open ocean (1800 m depth) summer in the Gulf of Mexico, suggesting summer (Johnson, 1978). Juveniles of this species are frequently spawning. Johnson (1978) found hydrated females in JuIy observed in symbiotic association with moon jeIIyftsh in the westem Gulf of Mexico. Ou the other hand, Aurelia aurita (Tolley, 1987). ln the SEBB C. ehrysurus is Flores-Coto & Sanchez-Ramirez (1989) found the S. a common species in littoraI zone, bay and setapinnis larvae all year-round with higher density during (Menezes & Figueiredo, 1980). spring-summer in the southern Gulf of Mexico. Typical coastal distribution was the characteristics of Distribution of S. vomer is limited in the westem C. ehrysurus larvae, most of them occurring in coastal water Atlantic and extends from Maine, USA, to Uruguay, living less than 30 m depth. This coincides with the results in shallow coastal waters. ln the SEBB, S. vomer larvae observed in the Gulf of Mexico (Leak, 1981; Flores-Coto were rare and their distribution was more restrlcted than & Sanchez-Ramirez, 1989; Shaw & Drullinger, 1990). that of S. setapinnis. Most of the larvae were found in Along the northwestern coast of a similar shallow waters (less than 40 m) from December to distribution was observed by Conand & Franqueville February, suggesting short and limited spawning during (1973), having a main distribution in depth zone of 20-50 summer. m. ln southem Brazil Weiss et aL, (1976) observed a different distribution pattern, i.e. they occurred from coast to 200 m isobath over the continental shelf along the Rio Grande do Sul State. Hydrographic conditions and larval distribution Larvae of C. ehrysurus were found alI year-round with major occurrences in December and J anuary in the SEBB, meanwhile, the peak of larval occurrences was in Oceanographic structure of this area has beeo August-November in the Rio Grande do Sul State, extensively studied since the pioneering work of Emílssoo southem Brazil (Weiss et ai., 1976). ln the eastem central (1961). There are at least tbree important water masses in AtIantic they were found mainly in August-November the southeastem Brazil, oamely, 1) Tropical water (TW) 74 Bolm Inst. oceanogr., S Paulo, 40(112), 1992

(Brazil Current), 2) South Atlantic Central Water sardine (Sardinops sagax). The reproductive strategy of (SACW), and 3) Coastal water (CW). Mixture of three the former during summer was attributed to the foUowing water masses through the heating process by solar factors: 1) enrichment of coastal waters by injection of radiation results in the formation of shelf water (Emílsson, nutrient rich SACW by upwelling process, 2) vertical op. cit.). stability of the water column, resulting in a high density of Hydrographic configuration over the continental food organisms in midwater, and 3) retention process by shelf depends mainly on the intensity of penetration of the large gyral circulation within the Bight. Similar SACW in the bottom layer over the continental shelf and physicallbiological processes are wel1 known in the coastal water which are influenced by many factors, such Southem Califomia Bight where many smal1 pelagic fishes as dominant wind system and intensity and meander of the spawn (Parrishet ai., 1983). Different seasonal and spatial Brazil Current. Remarkable seasonal variation was distribution pattems of larvae CaD be also observed for observed in the intensity of penetration ofthe SACW which carangid species. Some of them have restricted reached the coasta1 region during late-sprlng and summer occurrences to late- spring and summer, such as Oligoplites and retreated to offshore during winter (Matsuura, 1986). spp., Caranx spp., S. setapinnis and S. vomer larvae. On the Ascending process of the SACW in the coastal region by other hand, those showing year-around larvae occurrence the Elonan transport of surface CW offshore induces a have a preferential season, e.g. spring and summer for T. coastal upwelling (Miranda, 1982; Castro FIlho et al., 1987) lathami and summer for C. chtysurus and D. punctatus. which has been frequently observed at Cabo Frio and less This fact seems to demonstrates that most carangids frequently at Cabo Santa Marta Grande. This seasonal studied here preferred to spawn in austral summer using variation on hydrographic structure seems to have an the same strategy adapted by S. brasiliensis as observed by important influence on spawning season and area as well Bakun & Parrish (1990). as on the distribution pattem and abundance of fish larvae. When we compare the larval distributiODS of T. Since the original work on the aitical period concept lalham; andS. brasiliensis (Matsuura, 1983) we CaD observe proposed by Hjort (1914), many aUthors have investigated some similarity in distribution pattems during summer influence of environmental processes on reproductive season when both species occupy the entire continental success of fishes (e.g. May, 1974; Cushing, 1m; Loeb, shelf in the SEBB. Temperature and salinity variation in 1980; Lasker, 1975; 1978; Sinclair & Trembley, 1984). the positive statioDS for both species was similar, e.g. mean Meteorological and oceanographic conditions will temperature and salinity of the former was 23.74 °C (s = influence the availability of food organisIns and the eggs 355) and 34.95 psu (s = 0.81) respectively, and those of and larvae transport by residual current and thus will affect thelatterwere22.09°C(s = 2.67)and3S.31psu(s = 0.94). on reproductive success, resulting in recruitment variation Larvae of both species occur mainly in neritic regions, but (Norcross & Show, 1984). major concentrations of them on the same cruise didn't It is well known that most of the species have overlap. characteristic reproductive strategies, adjusting their However, we must consider differences in reproductive pattems with environmental factors, such as geographical habitats of the two species. S. bras;liensis in surface current. For example, in the northeast PaciflC the southwestemAtlantic is a typical tropical species which many coastal species which have pelagic stages of larvae occupies only the southeastem Brazilian coast (22 OS-30 OS) tend to spawn during winter, when wind-induced surface with relatively large population biomass. On the other transport directs onshore and not during the intensive hand, T. latham; inhabits a larger geographical area from upwelling season when there is a strong offshore transporto Cabo São Tomé (22OS) to Mar deI Plata (400S) in the ln the strong upwelling region in the northem California southwestem Atlantic and its spawning takes place in both whieh is characterized by strong surface Ekman transport temperate waters (10-11°C) (Cousseau, 1967) and tropical in the offshore direction, spawning of only small number of waters with surface temperature 27 SC, as shown in this surface spawning species takes place, avoiding quick reporto transport of their offspring into less productive offshore The largest occurrences of T. fathami larvae were region (parrish et ai., 1981). found on the cruise of Sep-Oct. 1976, meanwhile, only ln the southeastem Brazil it is not yet known the three S. brasiliensis larvae were found on the same cruise. details of the periodicity and amplitudes of these physical This suggests that spawning of T. lalhami takes place in processes over the biological process of the region and early spring, coinciding with the beginning of intrusion of reproductive success of fishes, but some interpretation CaD the SACW over the continental shelf, and extends into the be made based on the results obtained in other areas. summer. ln contrast, the spawning of S. brasiliensis takes Bakun & Parrish (1990) conducted a comparative study on place intensively in late-spring and summer. As can be reproductive habitats of two species of sardines in the seen in the commercial catches of the two species in the SEBB and California Current regions using the results southeastem Brazil, the population biomass of the S. obtained from the maritime meteorological data. They brasiliensis seems to be much larger than that of T.lathami. observed a considerable similarity in relation to The largest larval densities of the two species confirm this environment/life cycle between the two populations : observation, i.e. more than 1000 làrvae m·2 of S. brasiliensis Brazilian sardine (Sardinella brasiliensis) and Califomia (Matsuura, 1983) versus 108.4 larva m-2 of T. lalhami. KATSURAGAWA & MATSUURA: Carangid larvae: Distribution and abundance: Brazilian Bight 75

D. punctatus is another species which occupies References neritic regions and seems to apply the same reproductive strategy of the two species mentioned above, but the larval ABOUSSOUAN, A 1968. Oeufs et larves de abundance was much lower than that of T. lathami. téléostéens de 1'0uest africain. VI. Larves de C. chrysurus shows peak spawning in austral summer, Chloroscombrus chrysurus (Linné) et de Blepluuis but its distribution is basica11y related with the Coastal crinitus (Mitchill). (Carangidae). Buli. Inst. Water mass. Spawning strategy of this species is similar to fondament. Afr. noire, (sér. A), 30(3):226-237. that observed in with Harengula jaguana (another species of Oupeidae). H. jaguana eggs were col1ected exclusively APRIETO, V. L. 1973. Early development of carangid in coastal region (less than 37 km from the coast), but the fishes of the GuIf of Mexicoand the South Atlantic larvae showed a wider distribution (Matsuura, 1983). ln coast ofthe United States. PhD. Thesis. University both cases, after spawning part of larvae should be of Miami, Coral Gables, 179p. dispersed offshore with the Coastal Waters by Ekman transporto BAKUN, A & PARRISH, R. H. 1990. Comparative studies of coastal pelagic fish reproductive habitats: the Brazilian sardine (Sardinella aurita). J. Cons. int Explor. Mer, 46:'lfj9- 283.

BANNIKOV, A. F. 1987. On the , Resumo composition and origin of the Family Carangidae. J. Ichthyol.,27(1):1-8. No presente trabalho estudaram-se a distribuição e a abundância de larvas de peixes da famflia Carangidae da BAlLEY, K. 1989. Description and surface costa sudeste do Brasil, entre Cabo Frio (23OS) e Cabo de distribution of juvenile Peruvian jack mackerel, Santa Marta Grande (29"S). As amostras foram coletadas Trachun4s murphyi Nichols from the Subtropical com rede Bongô (malhagens de 0,505 mm e 0,333mm) em Convergence Zone of the central South PacifiCo arrastos oblíquos, durante dez cruzeiros oceanográficos Fishery Buli. natn. mar. Fish. Serv., U .S., com o N/Oc. "Prof.W.Besnard", do Instituto 87:273-278. Oceanográfico da USP e com N/Oe. "Cruz deI Sur", da PDP-SUDEPE. BERRY, F. H. & COHEN, L. 1974. Synopsis of the Foram identificados cinco táxons ao nível de espécie species of Trachurus (pisces: Carangidae). Quart. (Trachurus lathami, Chloroscombrus chrysurus, Decapterus J. FlaAcad. Sci., 35(4):1n-211. punctatus, Selene setapinnis e Selene vomer), além de quatro ao nível de gênero (Oligoplites, Caranx, Seriola e BERRY, F. H. & SMITH-VANIZ, W. F. 1978. Trachinotus). A espécie mais abundante foi T. lathami Carangidae. ln: FISHER, W. ed. FAO species (58,44% do total de larvas da familia Carangidae), seguida identification sheets for purposes. por C. chrysurus (15,22%) e D. punctatus (12,17%). T. Westem Central Atlantic. F'tshing Area 31. Rome, lolhami e D. punctatus apresentaram distribuição ampla FAO. v.1. por. toda a região nerítica, enquanto que C. chrysurus restringiu-se apenas às regiões mais próximas da costa. As CASTRO FlUlO, B. M.; MIRANDA, L. B. & larvas de carangídeos foram encontradas durante todas as MIYAO, S. Y. 1987. Condições hidrográficas na épocas do ano, mas na primavera e no verão observou-se plataforma continental ao largo de Ubatuba: a maior abundância, o que caracteriza maior intensidade variações sazonais e em média escala. Bolm Inst. de desova durante estas estações. Esse período de pico de oceanogr., S Paulo, 35(2):135-151. desova relaciona-se, aparentemente, à variação da estrutura hidrográfica da região. CIECHOMSKI, J. D. de. 1981. Ictioplancton. ln: BOLTOVSKOY, D. ed. Atlas deI zooplancton deI Atlantico Sudoccidental y métodos de trabajo con zooplancton marino. Contrib. Inst. naco Investig. Desarrollo pesq. (INIDEP),365:829-860. Acknowledgments ~_-:---;-:-_~_ & CASSIA, M. C. 1980. This research was funded by the FINEP Reproducción y fecundidad deI surel (Trachun4S (FInanciadora de Estudos e Projetos) and supported by the picturaÚls australis). Revta Investnes Desarrollo Oceanographic Institute of University of São Paulo. The pesq., 2(2):39-45. authors received the research fellowship of the CNPq (Conselho Nacional de Desenvolvimento Científico e ______& WEISS, G. 1973. Tecnológico) during this study. We thank Kazuko Suzuki Reproduccioén, desarrollo embrionario y larval deI and other laboratory technitians for technical assistance sure!, Trachun4s picturatus australis (Pisces) en el and drawing of figures. We also thank an anonymous area de Mar deI Plata y zonás adyacentes. Physis, A, reviewer for reviews and useful comments. B. Aires, 32(84):85-93. 76 Bolm Inst. oceanogr., S Paulo, 40(112), 1992

CONAND, F. & FRANQUEVILLE, C. 1973. GINSBURG, I. 1952. Fishes of the family Identification and seasonal distribution of carangid Carangidae of the northem Gulf of Mexico and larvae off the of Senegal and Gambia. Bun. three related speaes. Publs InsL mar. Sei., Univ. Inst. fondamenL Afr. noire, (sér. A), Sei. nat., Texas, 2:47-117. 35(4):951-978. GUSHIKEN, . S. 1988. Phylogenetic relationships of the Perciform genera of the family Carangidae. COUSSEAU, M. B. 1967. Contribuei6n aI Japan. J. Ichthyol, 34(4):8-461. conocimiento de la biología dei surel (Trachurus picturalis australis) dei área de Mar dei Plata HEMPEL, G. 1973. On the use or ichthyoplankton (Pisces, fam. Carangidae). Bom Inst. Biol mar., Mar surveys. FAO F"lSh. tech. Pap., 122:1-2. dei Plata, 15:1-38. HILDEBRAND, S. F. & CABLE, L. E. 1930. CUNHA, L. P. R. 1987. Importância da zona de Development and life history of fourteen arrebentação de praias para o desenvolvimento de teleostean fishes at Beaufort, N.C. Bun. Bur. juvenis de Trachinotus (Pisces, Carangidae): F"lSh., Wash., (46):453-464. aspectos da bioecologia e distribuição geográfica do gênero, com ênfase às espécies que ocorrem no HJORT, J. 1914. Fluctuation in the great fisheries litoral suVsudeste do Brasil e no Atlântico ocidental. of northem viewed ln the light of biological Tese de doutorado. Universidade de São Paulo, research. Rapp P.-v. Réun. Cons. perm. inL Explor. Instituto de Biociências. 146p. Mer, 20:1-228. .

CUSHING, D. H. 1977. Marine ecology and JOHNSON, G. D. 1978. Developmcnt of the fisheries. Cambridge. Cambridge University Press. fishes of the mid-Atlantic Bight. AD atlas of 278p. egg. larval and juveniJc stages. vol IV. Carangidae through Ephippidae. U .S. Fish Wildl. Servo DITTY, J. G. & TRUESDALE, F. M. 1984. Program FWS/OBS-78-U. 314p. Ichthyoplankton surveys of nearshore Gulf waters off Louisiana: January-February and July KATSURAGAWA, M. 1990. Estudo sobre o 1976. ASB Bun., 31(2):55-56. desenvolvimento, a distribuição e a abundâneia de larvas de carangfdeos da costa sudeste do EMÍLSSON, I. 1961. The shelf and coastal waters Brasil. Tese de doutorado. Universidade de off southem Brazil. Bolm Inst. oceanogr., S Paulo, São Paulo, Instituto Oceanográfico. 155p. 11(2):101-1U. KRAMER D.; KALIN, M. J.; STEVENS, E. G.; FAHAY, M. P. 1975. AD annotated list of larval TIlRAILKILL, J. R. & ZWEIFEL, J. R. 1972. and juvenile fishes captured with a surface-towed Collecting and processing data on fish eggs "and meter net in the South Atlantic Bight during four RV larvae in the Califomia Current region. U. S. Dept. Dolphin cruises between May 1967 and February of Commerce. Circo natn. mar. F"1sh. Serv., U.S., 1978. Spec. seient. Rept natn. mar. F"lSh. Serv., U.S., (370):1-38. F"1sheries, 685, 39p. LASKER, R. 1975. Field criteria for survival of anchovy larvae: the relation between inshore FAO. 1989. Fishery statistics: catches and landings chlorophyl maximum layers and successful first 1987. Yb. fish. StatisL, 64:1-490 .. feeding. Fishery Bull. natn. mar. F"1sh. Serv., U.S., 73(3):453- 462. FARRIS, D. A. 1961. Abundance and distribution or eggs and larvae and survival of larvae of jack ______. 1978. The relationship between mackerel (Trachurus symmetricus). Fishery Bu11. oceanographic conditions and larval anchovy food natn. mar. Fish. Serv., U.S., 61(187):247-279. in the California Current: Identification of factors contributing to recruitment failure. Rapp. P-v. FWRES-COTO, C. & SANCHEZ-RAMIREZ, M. Réun. Cons. perm. int. Explor. Mer, 173:2U- 230. 1989. Larval distribution and abundance of Carangidae (PISCES), from the Southem Gulf or LEAK, J. C. 1981. Distribution and abundance Mexico.1983-1984. GulfRes. Repts, 8(2):117-128. of carangid flSh larvae in the eastem Gulf of Mexico 1971-1974. BioL Oceanogr., 1:1-28.

FUKUHARA, O. 1987. Recent trends in maricu1ture LOEB, V. L. 1980 Patterns or spatial and species and seed production of fish in southem Japan. abundance within the larval fish assemblage of the Tech. RepL natn. mar. Fish. Serv., U.S., 47:1-2. north Pacifi~ Central gyre duringlate summer. Mar. Biol,60:189-200. KATSURAGAWA & MATSUURA: Carangid larvae: Distribution and abundance: Brazilian Bight 77

RICHARDS, E. J.1985. Status ofthe identification ofthe MATSUURA, Y. 1983. Estudo comparativo das fases early life stages of fishes. Bull. mar. ScL, iniciais do ciclo de vida da sardinha verdadeira. 37(2):756-760. SardineOa brasiliensis e da sardinha cascuda, . Harengula jaguana (Pisces: Clupeidae) e nota sobre SACCARDO, S. A 1980. Biologia e bionomia de a dinâmica da população de sardinha verdadeira na Trachurus lathami Nichols, 1920 (Teleostei: região sudeste do Brasil. Tese de livre-docência. Carangidae) na plataforma continental brasileira Universidade de São Paulo, Instituto Oceanográfico. entre 230S (RI) e 3

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