432 I <#*>" °' c o, ^7-ES 0< »■ NOAA Technical Report NMFS Circular 432 Synopsis of Biological Data on Bonitos of the Genus Sarda May 1980 ■I t/1 \s> o n to C r i_ (1) 1- a u CO c/) CO (/) 11 Figure 3. — Distribution of the fertilized eggs of Sarda chiliensis lineolata off Baja California in the spring of 1966, based on tows with the ichthyoplankton net. 1 — from 1 to 50; 2 — from 51 to 500; 3 — more than 500 in one tow of the egg net; 4 — ichthyoplanktonic stations; 5 — isotherms. (From Sokolovskii 1971.) distribution of the fertilized eggs of S. orientalis. Mito (1961) gave generalized descriptions of fertilized eggs of the species in what he termed the Scombrina, including S. orientalis. However, Mito gave no capture locality for his samples. Except for this fragmentary bit of informa- tion on the planktonic fertilized eggs of S. orientalis off Japan, there is nothing on the distribution of their eggs in the other areas of the Pacific and Indo-Pacific where this species occurs. Records of larval S. orientalis are few. Gorbunova (1963) recorded the capture of a larva 11.56 mm long from off the northwest coast of Australia in the Indian Ocean. Elsewhere in the Indian Ocean, Jones (1960) re- ported specimens 80, 89, 174, and 262 mm in total length from Vizhingam on the west coast of India. In the Paci- fic Ocean, juveniles ranging in length between 75 and 100 mm are reportedly caught in waters adjacent to south- ern Kyushu (Yabe et al. 1953), and Kishinouye (1923) re- ported a specimen 170 mm taken off Wakayama Pre- fecture, Japan. Juveniles from the stomachs of predators have been recorded from Hawaii (Honolulu Laboratory, Southwest Fisheries Center, unpubl. data.) Yabe et al. (1953) reported that 75-100 mm S. orien- talis are caught in waters around southern Kyushu. They also reported on S. orientalis landed at the port of Aburatsu, Japan, ranging in size from 161 to 348 mm. Preadult S. orientalis occasionally appear in large numbers in the Indian Ocean. Jones (1960) reported large catches offish ranging in length from 160 to 200 mm off the west coast of India between Trivandrum and Cape Comorin. Gnanamuttu (1966) recorded the occurrence of S. orientalis ranging from 240 to 260 mm in total length off the northeast coast of India. In the waters around Sri Lanka the commercial fishery lands fish ranging up- wards from 20 cm (Sivasubramaniam 1969). include two larvae 2.9 and 3.5 mm long; Klawe (1961b) could not determine whether they were S. chiliensis or S. orientalis. Other than the prelarval stages which Barn- hart (1927) and Orton (1953a, 1953b) reported, and the two larvae recorded by Klawe, the only other larvae of Sarda reported from the eastern Pacific are those by Sokolovskii (1971). Sokolovskii reported the capture of six larvae ranging from 7 to 13.8 mm at an ichthyo- plankton station in the waters off Baja California. It can be seen from Table 7 that juvenile S. chiliensis in the Northern Hemisphere have been found off Baja Cali- fornia between lat. 22° and 26° N. In the Southern Hem- isphere they have been found along the coast of Chile and Peru between lat. 13° and 25°S. Vildoso (1966) sampled preadult S. c. chiliensis from the Peruvian fishery rang- ing upward from 20 cm, and Barrett (1971) reported bonito of about 15 cm taken with anchovy catches at Arica, Chile. In southern California, preadult S. c. lineo- lata 15.2-25.4 cm (6-10 in) are first seen by the bait fishermen in the early summer (Frey 1971). Sarda orientalis There is hardly any information in the literature on the Sarda sarda The distribution of fertilized eggs of S. sarda has been studied in greater detail in the Black Sea and the Sea of Marmara. Mayorova and Tkacheva (1959) reported on the distribution of S. sarda eggs in the Black Sea in 1956 and 1957 (Fig. 4). They noted that in 1956 bonito eggs were found in great numbers in the surface layer, the maximum catch amounting to 12,000 eggs per "10-min catch," presumably made using a plankton net. At a depth of 5 m the number of eggs taken did not exceed 5,000 eggs/10-min tow. In 1957 the overall abundance of fertilized eggs was less than in 1956, and the vertical distribution was also different in that the 5 m depth pro- duced more eggs than the surface layer. Demir (1963) noted that available data indicate that S. sarda eggs have been found everywhere in the Black Sea, the loca- tions varying from year to year. The fertilized eggs of S. sarda have also been found in the Sea of Marmara (Fig. 5) (Demir 1963). It can be seen from Figure 5 that in June and July of 1959 most of the fertilized eggs were found in the eastern portion of the Sea of Marmara. 12 Table 7. — Records of capture of juvenile Sarda chiliensis from the eastern Pacific. 1961b, table 2, 1962.) (From Pinkas 1961, table 2; Klawe Date General locality Latitude Longitude Method of capture Size range Fork length (mm) No. of speci- Northern Hemisphere 17-18 May 1947 Off La Jolla, Calif., United Night light, — 1 States dip net 5 Aug. 1951 100 miles NW of Cape San Lazaro Baja Calif., Mexico 25°35'N 113°56'W Night light, dip net 42 1 5 Aug. 1951 Off Baja Calif., Mexico 25°35'N 113°56'W Night light, dip net 42 1 12 Aug. 1951 Off Ballenas Bay, Baja Calif., Mexico 26°29.5'N 133°29.2'W Plankton net 2.9,3.5 l 1 15 July 1953 South of Cape San Lazaro, Baja Calif., Mexico 23°47'N 112°25'W Night light, dip net 25-41 5 18 July 1953 Southwest of Cape San Lazaro, Baja Calif., Mexico 23°16'N 112°45'W Night light, dip net 19-41 8 11 Apr. 1955 South of Cape San Luca9, Baja Calif., Mexico 22°52.8'N 109°53.7'W Night light, dip net 16.7 1 11 July 1956 Southwest of Cape San Lucas, Baja Calif., Mexico 23°35'N 112°11'W Night light, dip net 33.0-33.5 2 12 July 1956 Southwest of Cape San Lucas, Baja Calif., Mexico 22°20'N 112°27'W Night light, dip net 24-48 8 17 July 1956 Southwest of Cape San Lucas, Baja Calif., Mexico 22°47'N 112°14'W Night light, dip net 54.5 1 5 Apr. 1960 Off Baja Calif., Mexico 23°19'N 110°22'W High speed net 92 1 Southern Hemisphere 20 Feb. 1951 OffPt. Lobos, Peru ? 125-130 3 3 Jan. 1956 OffPt. Negra, Peru Bait net 143-164 3 10 Dec. 1957 Independencia Bay, Peru 14°14'S 76°12'W Bait net 128 1 13 Dec. 1957 Off Ilo, Peru 17°38'S 71°23'W Bait net 121 1 15 Dec. 1957 Off Ilo, Peru 17°38'S 71°18'W Bait net 200.9 1 18 Dec. 1957 Off Ilo, Peru Bait net 70-135 3 21 Dec. 1957 South of Santa, Peru 09°31'S 78°26'W Bait net 38.0-51.0 5 31 Dec. 1957 OffPt. Pichalo, Chile 19°35'S 70°16'W Night light, dip net 37 1 1 Jan. 1958 Off Ilo, Peru 17°47'S 71°30'W Bait net 86.8 1 3 Jan. 1958 OffPt. Dos Reyes, Chile 24°30'S 70°49'W Night light, dip net 39 1 3 Jan. 1958 Off Pt. Dos Reyes, Chile 24°36'S 71 o 01'W Night light, dip net 36-44 3 14 Jan. 1958 Southwest of Fraile Pt., Peru 13°14.8'S 77°55.5'W Night light, dip net 34 1 Feb. 1958 Off Chimbote, Peru Bait net 89-129 3 6 May 1958 Almejas Bay, Baja Calif., Mexico Bait net 103 1 1 Feb. 1959 Sama Cove, Peru Bait net 111 1 15 Mar. 1959 Off Ilo, Peru ? 135-160 3 16 Mar. 1959 Off Barranca, Peru ? 173, 199 2 Identity uncertain. It is of interest that very little is known of the distri- bution of fertilized S. sarda eggs in the Mediterranean Sea. Demir and Demir (1961) noted that planktonic fer- tilized eggs of S. sarda were unknown from the Medi- terranean Sea. They pointed out that the eggs reported on by Sanzo (1932) were eggs that were extracted from a female gonad and artificially fertilized and reared. More recently, however, Duclerc et al. (1973) reported on the collection of fertilized S. sarda eggs near the Balearic Islands in the Mediterranean Sea. Thus, as far as can be determined from the literature, the distribution of ferti- lized eggs of bonito, is not very well defined in the Medi- terranean Sea. Except for the report of the collection of the fertilized eggs of S. sarda off the coast of Massachusetts by Sette (1943), nothing is known of their distribution in the Atlantic Ocean. Mayorova and Tkacheva (1959) stated that S. sarda prelarvae and larvae occur in the same regions as the fertilized eggs in the Black Sea. Figure 4 shows that fer- tilized eggs of S. sarda are widely distributed in the Black Sea, thus indicating that larval bonito are also 13 Egg-net catches (number per 10 minutes) EH) ot to AO 100 KM— 200 2(H— 500 5W— 1000 m— zoon O over 2000 CD not found - II- 50 - 51-100 - iui-200 - c'ul-JOO - }OI-5oO - 5uI-lOuO Figure 4.— Distribution of the fertilized eggs of Sarda sarda in the Black Sea, in June 1956 (a) and June 1957 (b). (From Mayor- ova and Tkaeheva 1959, figs. 1, 2.) widely distributed. The larvae were more abundant at a depth of 5 m than at the surface. Mayorova and Tkaeheva also noted that juvenile S. sarda 10-95 mm long were distributed over a vast area of the Black Sea in waters from 80 to 100 miles from the coast. In the Sea of Marmara, however, plankton tows did not reveal any S. sarda larvae although fertilized eggs in all stages of de- velopment were plentiful (Demir 1963). Elsewhere, larvae of 5. sarda have been reported from the Mediterranean Sea northwest of Oran by Ehren- baum (1924) and in the Alboran and Mediterranean Seas by Buen (1930, 1932). More recently, freshly hatched 14 CO _l ^ c ^ x 5 w - 1 " < £E Z -I =) I— Q_ V iLl < z z o i ■ cr Q UJ UJ -^ $ zj -, co ~3 £: UJ QSz 2 tr uj 5 = 1 UJ O ^ CC 2 (E (0 O 3 o -1 O O O UJ °o CO o O t\J O 1 u o i 2 uj UJ UJ UJ cc pc X UJ UJ > £ £ CO g g I O O t- p p o CO CO _l o • © 6 u i 10 bl IP 15 prelarvae were collected near the Balearic Islands by Duclerc et al. (1973). In the Atlantic Ocean, larvae have been found off Cuba (Gorbunova and Salabarria 1967), and a juvenile measuring 34 mm has been recorded off the South Carolina coast (Klawe 1961a). Two juvenile S. sarda 64 and 67 mm in total length were also captured in the Gulf of Mexico (Klawe and Shimada 1959). Sarda sarda from 6 to 18 cm long are captured in vary- ing numbers in the Dardanelles, the Bosporus, the Sea of Marmara, and the Black Sea nearly every year in July- August (Demir 1963). The abundance of preadults in this size range apparently is a good indicator of the future abundance of fish 25-38 cm long which are the basis of an important Turkish fishery that takes place starting in September in the same areas. Preadults 8-12 cm long have been captured in large numbers in the Aegean Sea (Serbetis 1955), and Belloc (1954) discussed the occur- rence of S. sarda 7.5-20 cm long caught in the Gulf of Naples and the Gulf of Catania and 15-25 cm long fish in the Bay of Biscay. The capture of preadults 20-30 cm long has been re- ported from the Atlantic coast of France (De la Tourrasse 1957), and 25-35 cm fish have been taken commercially together with mackerel in the Miramichi estuary in the Gulf of St. Lawrence (McKenzie 1959). Klawe and Shimada (1959) reported the capture of two specimens 64 and 67 mm long in the Gulf of Mexico. Young S. sarda 12.7-15.2 cm (5-6 in) long have been taken off Orient, N.Y. (Nichols and Breder 1927). 2.22 Adults Sarda australis Information on the seasonal and annual variations in the occurrence of S. australis on the east coast of Australia is fragmentary. Serventy (1941b) stated that this species apparently is only a summer visitor to Vic- toria. Munro (1958) noted that S. australis was more plentiful north of Sydney in the winter; and according to Grant (1972), it commonly occurs in schools in inshore waters of Queensland, especially in winter. However, Marshall (1964) stated that S. australis is very common on the coasts of Queensland and New South Wales and is found in great schools throughout the year. Sarda chiliensis For the northeast population of S. chiliensis MacCall et al. (1976) suggested an uneven geographical distribu- tion of various age groups. They noted that, roughly, older fish were more available offshore and around Mex- ico although large fish were also taken in the Santa Bar- bara, Calif., area in the fall. As for the southeast population of 5. chiliensis, An- cieta (1963) found that off the coast of Peru 82.2% of the total landings during 1951-60 were made in the central zone, from Chimbote to Pisco; 11.8% were made in the southern zone, from San Juan to Ilo; and 6% were made in the northern zone, from Mancora to Salaverry. How- ever, Mejia et al. (1971) 3 noted that beginning in 1962 there has been a gradual overall decline in the annual Peruvian landings of 5. c. chiliensis, most notably at the central zone port of Callao; in contrast the landings at the northern zone port of Paita increased somewhat. Off the Chilean coast S. c. chiliensis are taken from the northern boundary of Chile to as far south as Talca- huano, but most are taken off the northern ports of Arica, Iquique, and Antofagasta, especially between Iquique and Mejillones (Barrett 1971). It was also seen that almost two-thirds of the annual Chilean landings of S. c. chiliensis are made in the last third of the year. Sarda orientalis Along the southwest coast of India, fully mature adult S. orientalis are found from May to September followed by the juveniles from October to November. Slight an- nual fluctuations in seasonal occurrence are also seen (Silas 1964). Off the south coast of Sri Lanka (Ceylon), S. ori- entalis appears in the commercial catches throughout the year. Mature fish are taken off the south and south- west coasts primarily between September and February, and the juveniles appear off the west coast more often from June to August (Sivasubramaniam 1969). Gen- erally, S. orientalis appears in the commercial catches most frequently off the south coast, to a lesser extent off the west and northwest coasts, and least frequently off the southwest and northeast coasts. Sarda sarda The seasonal occurrence of S. sarda in various locali- ties is summarized in Tables 8 and 9. 2.3 Determinants of distribution changes Along the California coast there appears to be a re- lationship between the latitudinal distribution of S. chiliensis and sea temperature (Radovich 1963). An up- surge in the average daily S. chiliensis catch of party- boat anglers took place with the onset of a warm water pe- riod off California in 1957. Radovich suggested that S. chiliensis moved northward into California waters during warm years. He also noted that during warm years, bonitos spawned successfully in California waters, thus further increasing the local population. From all indications it appears that all the species of Sarda are closely associated with the coast, where the ocean environment may be subject to rapid changes. Demir (1963) stated, however, that S. sarda cannot toler- ate sudden changes in environment, but they can adapt to gradual changes in temperature ranging from 12° to 'Mejia, J., M. Samame, and A. de Vildoso. 1971. Revision de la pesqueria del bonito en quas Peruanas. Fourth Session of FAO Panel of Experts for the Facilitation of Tuna Research, La Jolla, Calif., 8-13 November 1971. (Mimeogr.) 16 Table 8. — Fishing seasons for Sarda sarda. Area Fishing season Peak months of fishing Source Black Sea Bosporus, Sea of Marmara, Dardanelles Aegean Sea (around Greece) Eastern Mediterranean (off Spain) Mediterranean (off Tunisia) Eastern tropical Atlantic Eastern Atlantic (off Morocco) Eastern Atlantic (Bay of Biscay) Western Atlantic (Gulf of Maine) May-October Throughout the year Throughout the year May-June Throughout the year October-May Throughout the year Mid-April to mid-May June-October April-May, September-December May-July Demir(1963) Demir (1963) Serbetis (1955) Gruvel (1931) Postel (1955b) Postel (1955b) Furnestin et al. (1958) delaTourrasse(1957) Bigelow and Schroeder (1953) Table 9. — Fishing seasons in the Spanish fishery for Sarda sarda. (From Rodriguez-Roda 1966, table 2.) Area Fishing season Peak months of fishing Bay of Biscay (along Spanish coast) Eastern Atlantic (off northwest coast of Spain) Eastern Atlantic (southwest coast of Spain) Western Mediterranean (southeast coast of Spain) Western Mediterranean (east coast of Spain ) Western Mediterranean (northeast coast of Spain) Western Mediterranean (Balearic Islands) Eastern Atlantic (Canary Islands) April-November April-December January -December January-December January-December January-December January-December January-December July-October July, August September, October August-October May, September June-November May-September August, September to identify the sex of another bonito except by beha- vioral characteristics. The bonito is normally hetero- sexual; however, Vildoso (1960) found cases of hermaph- roditism in S. c. chiliensis in Peruvian waters. It is of interest that Magnuson and Prescott observed that one fish exhibited both a "wobbling" and "following" beha- vior. They attributed the anomalous behavior of this particular fish to a possible misidentification and did not speculate on the possibility of hermaphroditism or other causes for the aberrant behavior. As is the case with the other species of bonitos, the sexes cannot be distinguished in S. sarda. Apparently, however, some Turkish fishermen can distinguish the sexes by rubbing the skin anterior of the anus with a fingertip. The skin is said to be smooth in the females and rough like emery paper in the males (Demir 1963). 3.12 Maturity Sarda australis 27 °C and salinities between 14 and 39%o. Sarda sarda have been taken in the Miramichi estuary, Gulf of St. Lawrence, where the condition must surely be less than oceanic. Sarda orientalis have been found in waters rang- ing from 13.5° to 23°C (Kishinouye 1923). 3 BIONOMICS AND LIFE HISTORY 3.1 Reproduction 3.11 Sexuality All four species of Sarda are usually heterosexual, and there are no apparent external anatomical differences be- tween the males and females. In Sarda c. lineolata, how- ever, Magnuson and Prescott (1966) noted a sexual di- morphism in the behavior of this species in captivity. Magnuson and Prescott noted that some S. c. lineolata in their observation tank were "wobblers" and that others were "followers." They postulated and subsequently de- termined that the "wobblers" were females and the "fol- lowers" were males. They also determined by their observations that a male bonito was apparently unable The literature contains very little information on Sarda australis. Serventy (1941b) reported on three fish caught in January and February 1939 off Wilson's Promontory and Port Albert. The fish weighed between 1.8 and 2.3 kg (4 and 5 lb); and two, which were females, had "relatively large roes." Whitley (1964) noted that S. australis sometimes venture into Victoria in the summer (January to April) and that some of these fish had large roe. He also found that fish ready to spawn were found in February and March off New South Wales. Sarda chiliensis Barrett (1971) determined the gonad index (GI) (GI = t^tX 10 2 , where w weight of both ovaries in grams and W = body weight in grams) of bonitos and found that female S. c. chiliensis off the coast of Chile initially reached sexual maturity at a length of 51 cm. For the bonitos off the coast of Peru, Vildoso (1966) determined that the size at first spawning ranged from 47 to 53 cm. Kuo (1970) stated that in the Northern Hemisphere population of S. chiliensis, the females attained sexual maturity at 51 cm and that the fish was 5 yr old at that length. 17 Sarda orientalis Although the size of S. orientalis at first spawning has not been clearly determined, a 38.6 cm specimen posses- sed residual eggs which were presumably remnants from an earlier spawning (Silas 1964). However, Silas indi- cated that specimens in "ripe running condition or had already spawned, some showing signs of recovery" measured between 480 and 605 mm from samples col- lected in 1960 and 1961 at Vizhingam, India. The age of these specimens was not determined. Rao (1964) found four specimens ranging in size from 48 to 55 cm with running-ripe ovaries in June, August, and September 1959, also from Vizhingam. Sarda sarda The bonito in the Sea of Marmara and the Bosporus usually attain sexual maturity at the end of the second year, although in some years they may be sexually ma- ture in a year. Two-year-old fish range from 52 to 57 cm and 1-yr-old fish from 42 to 48 cm. In the Black Sea, part of the stock of S. sarda attains sexual maturity in the second year of life. These 2-yr-old fish are said to vary from 33 to 50 cm. Off the coast of Dakar in the eastern Atlantic, the size at first maturity is 392 mm for the males and 370 mm for the females. These fish are less than a year in age. 3.13 Mating As Magnuson and Prescott (1966) pointed out, pelagic schooling fish were assumed to have no discrete court- ship behavior and no pairing and were believed to shed eggs and sperm promiscuously while gathered in large ac- tive schools. To the contrary, Magnuson and Prescott found that S. c. lineolata do exhibit courting and pairing behavior, if only temporarily, and observed pairs of fishes in a sequence of behavior leading to a simultaneous and adjacent release of eggs and milt. The pair of bonitos re- leased the gametes during a circle swimming behavior in which the male swam in tandem with the female in a circular path. Although some behavioral patterns have been observed only in 5. c. lineolata, it would not be unreason- able to expect that the other species of Sarda also be- have in a similar manner. 3.14 Fertilization External in all four species of Sarda. 3.15 Gonads Sarda australis No information. Table 10. — Fecundity of Sarda sarda. Size of Fecundity Area fish (No. of eggs) Reference Black Sea 40-50 cm 700,000-1,000,000 Zusser (1954) 60-70 cm 1,500,000-2,000,000 70 cm 6,000,000 Black Sea — 450,000-1,000,000 Slastenenko (1956) Black Sea — 700,000-1,000,000 Krotov (1957) Black Sea 56-65 cm 732,160-3,233,580 Mayorova and Tkacheva (2-3.8 kg) (1959) Eastern 60 cm 900,000 Postel (1955a) Atlantic Sarda chiliensis Vildoso (1963a) estimated the number of ova spawned by a 600 mm (3 kg) bonito at half a million per spawning season. According to Vildoso, spawning is fractionary. Kuo (1970) estimated that the fecundity of the northern S. chiliensis ranged from 104,900 to 894,200 eggs for fish from 47.6 to 63.7 cm in fork length. He indicated that the fecundity increased exponentially with size of fish. Sarda orientalis Silas (1964) made fecundity estimates for five mature specimens of S. orientalis from the Indian Ocean. He estimated that the females produce 0.08-0.15 million eggs/spawning and 0.24-0.64 million eggs/spawning season. Rao (1964) estimated that the bonito spawns 0.21-0.28 million eggs/spawning and 0.91-1.15 million eggs/spawning season in the Indian Ocean. Sarda sarda Demir (1963) summarized the fecundity determina- tions for S. sarda made by various investigators in the eastern Atlantic (Table 10). The fecundity estimates range from 450,000 to over 3,000,000 eggs. 3.16 Spawning Sarda australis Whitley (1964) indicated the occurrence of fish that were ready to spawn in February. Sarda chiliensis The spawning season for S. c. chiliensis in Chilean waters begins in September, is at a maximum in October and November, and is well over before April (Barrett 1971). The spawning season off the coast of Peru is very similar to that off Chile; peak spawning of bonito extends from October to February (Vildoso 1966). According to both authors, larger, older bonito mature earlier in the spawning season than do the younger fish (Fig. 6). For the Northern Hemisphere population of S. chiliensis, Kuo (1970) investigated several different ap- proaches to determine the spawning season, including 18 NOV. a APR.o o o SEPT. x' JULY o--' 46 48 50 52 54 56 58 60 62 64 66 68 LENGTH (cm) Figure (i. — Mean monthly gonad indices, GI = U' 10 , by length class, of female Sarda chiliensis chiliensis sampled from the com- mercial landings at Iquique, September 1968 to July 1%9. (From Barrett 1971, fig. 5.) the annual cycle in the development of intraovarian oocytes, the annual cycle of frequency distribution of ovum diameters, and the annual change in gonad index. All of the different methods indicated that the spawning season for the Northern Hemisphere S. chiliensis in southern California is from May through July. Kuo speculated that off southern Baja California spawning may start as early as April and may continue into August. He also indicated that the larger fish tend to spawn earlier in the season and longer than the smaller fish. Frey (1971), however, attributed an early spawning season to S. c. lineolata between southern California and northern Baja California waters. He stated that the bonitos spawned between January and May in these waters. The results of an ichthyoplankton survey off Baja Cali- fornia seem to corroborate the conclusions of Kuo (1970). On the basis of the distribution of fertilized eggs, Soko- lovskii (1971) showed that S. c. lineolata spawned in a relatively large area near the coast (Fig. 3). In early March the most intensive spawning occurred in the southern area and in early April it occurred in the north- ern area. The depth of the ocean in the spawning area ranged from 40 to 150 m, and practically no bonito eggs were found outside of the 200 m isobath. Larval bonito were captured at only one station during this survey. Most of these observations tend to support Klawe (1961b) who, on the basis of the capture of larval and ju- venile S. chiliensis, had stated that the spawning of this species takes place in the warmer season off California, Baja California, Peru, and northern Chile. Evidence has also been gathered which suggests that 1- and 2-yr-old S. c. lineolata spawn in areas influenced by warmwater discharges during spawning periods when the water is cold. Tag returns have shown that small numbers of young fish stay in these areas for as long as 2 yr and provide a small amount of recruitment even in cold years (Collins and MacCall 1977). Sarda orientalis The literature contains very little information on 5. orientalis spawning in the eastern Pacific and around Hawaii. Klawe (1961b) reported on two larval forms of Sarda from off Baja California that could not be identi- fied to species. The distribution of the northeast popula- tion of S. chiliensis overlaps or nearly overlaps that of the population of S. orientalis at the location of capture of the larvae; therefore, the larvae could be either of the two species. Nothing is known about the spawning of S. orientalis around Hawaii except that a few juveniles have been found in the stomachs of predators (Honolulu Labora- tory, Southwest Fisheries Center, unpubl. data), which provides evidence of reproduction. No formal studies on the spawning of S. orientalis in Japanese waters have been made. Kikawa and Staff of the Nankai Regional Fisheries Research Laboratory (1963) presumed that spawning occurs in the coastal waters of the tropical zone in the Lndo-Pacific. There ap- parently is some local unrecorded knowledge of bonito spawning in Japan because Harada et al. (1974) re- ported that the spawning season is May-June. They caught mature S. orientalis on 15 May and 10 June in traps set near the coast of Oshima Island and used these fish successfully in an artificial fertilization experiment. Mito (1961) gave descriptions of the fertilized eggs of S. orientalis found in plankton collections but gave no col- lection details. Kishinouye (1923) and Yabe et al. (1953) reported on the capture of juvenile bonito in southern Honshu and Kyushu waters. Off the coast of Vizhingam in the Indian Ocean, S. ori- entalis spawn from April to September and possibly in other months of the year. Ova diameter frequency distri- butions indicate the possibility that individual females spawn several batches of ova (Fig. 7) during the spawn- ing season (Silas 1964). Gorbunova (1963) reported on the larva of S. ori- entalis from the Indian Ocean off the northwest coast of Australia and generalized that spawning is limited to the autumn-winter period. However, this conclusion is based, apparently, on the capture of one larva of S. ori- entalis in December. Gorbunova probably has the seasons confused, for December off the northwest coast of Australia would be closer to summer in the Southern Hemisphere. Sarda sarda Demir (1963) stated that S. sarda spawning begins in mid-May, reaches a peak in June, and lasts at least to the end of July. He assumed that the spawning season is the same in the Black, Marmara, and Aegean Seas. Indications are that the spawning season in the Medi- 19 Table 12.— Relation between the stage of sexual maturity and the gonadosomatic index (GI =~tf * 100, where w is gonad weight and W is fish weight) for Sarda sarda landed at Barbate, Spain in 1964. (From Rodriguez-Roda 1966, table 26.) 10 20 30 40 50 60 OVA DIAMETER ( IN MICROMETER DIVISIONS ) 70 Figure 7. — Ova diameter-frequency polygons of ripe ovaries of the oriental bonito, Sarda orientalis, from the southwest coast of India. The ripe ova were sampled from two of four ripe fish examined from June to September 1959. (From Rao 1964, fig. 3.) terranean is also from May to July. However, Demir (1963) noted that Dieuzeide et al. (1955) found that the spawning season off the coast of Algeria is from March to May. Rodriguez-Roda (1966) made observations on the sex- ual development of S. sarda landed in May, June, and July 1963 and 1964 in the Spanish fishery based at the Mediterranean ports of Barbate and Tarifa. By gross examination of the gonads, he classified the males and females according to six stages of sexual maturity: I, Im- mature; II, Early maturing; III, Mature; IV, Pre- spawning; V, Spawning; and, VI, Spent. Data on the monthly percentage distribution of fish in the various stages of sexual maturity were given by Rodriguez-Roda (Table 11). He also computed the GI of the fish for his Males Females Stage of sexual No. of No. of maturity Mean Range fish Mean Range fish I (immature) 0.56 0.42-0.69 2 II (early maturing) 0.94 — 1 0.85 — 1 III (mature) 4.15 0.96-6.90 30 4.44 1.77-12.15 23 IV (prespawning) — — — 5.39 3.56-9.40 8 V (spawning) — — — — — — VI (spent) — — — 2.15 — 1 w 1964 samples using the equation GI = xy • 100, where w is the weight of the gonads and W is the weight of the fish, and determined the relation between the stage of sexual maturity and the GI (Table 12). Rodriguez-Roda noted that the smallest fish with prespawning and spawning gonads were 39.5 cm for the males and 40.5 cm for fe- males. In the eastern Atlantic near the coast of Dakar the spawning season extends from December to June, in- cluding peaks in January and April, and June to July in Moroccan waters. Less is known about S. sarda spawning in the western Atlantic. Bigelow and Schroeder (1953) stated that the bonito spawns south of the Gulf of Maine in June. However, Sette (1943) reported the collection of bonito eggs near Martha's Vineyard, Mass., in July. Further south off the South Carolina coast Klawe (1961a), based on the capture of a singe 34 mm bonito, stated that this species spawns in the winter. The cap- ture of a larval bonito (Gorbunova and Salabarna 1967) off the coast of Cuba and a juvenile specimen in the central Gulf of Mexico (Klawe and Shimada 1959) indi- cates some spawning activity in these areas. 3.17 Spawn The fertilized eggs of the bonitos have been described for all the species in the genus except Sarda australis. Except for differences in size, the fertilized eggs at the various developmental stages appear to be quite similar, and it is doubtful if the species could be separated. As ex- amples of differences in size of the fertilized eggs among the species of Sarda, Demir (1963) found that for S. sarda in the Sea of Marmara, the eggs, which were measured after preservation in 4% Formalin, varied between 1.18 and 1.55 mm in diameter. The fertilized eggs of S. c. Table 11. — Monthly percentage distribution of Sarda sarda in the various stages of sexual development, Barbate and Tarifa, Spain, in 1963 and 1964. (I, immature; II, early maturing; III, mature; IV, prespawning; V, spawning; VI, spent.) (From Rodriguez-Roda 1966, table 24.) Males Females No. of No. of I II III IV V VI fish I II III IV V VI fish May 7.02 1.75 87.72 3.51 57 30.00 15.00 55.00 — 20 June — — 73.81 23.81 2.38 — 42 2.38 — 64.29 30.95 — 2.38 42 July — — 100.00 — — 18 — — 54.17 41.67 — 4.17 24 20 Uneolata from off the coast of Baja California measured from 1.4 to 1.8 mm in diameter (Sokolovskii 1971). How- ever, no mention is made of whether the eggs were measured before or after preservation. Harada et al. (1974) reported that the fertilized eggs of S. orientalis were from 1.32 to 1.45 mm in diameter. Here again it is not clear whether the eggs were measured fresh or after preservation. These fertilized eggs were the result of a successful experiment in artificial fertilization, the first record of success in artificial fertilization for this species. Because the fertilized eggs of the species of Sarda are similar in appearance, as a typical example Demir's (1963) description of the eggs (Fig. 8) of S. sarda is given below. The fertilized planktonic eggs of S. sarda are spheri- cal, transparent, and, except for the oil globule, color- less. The yolk sac is homogeneous and finely granulated, and varying numbers of oil globules of different sizes are present on its surface. Although yellow and black pigmentation are present on the developing embryo, only the black pigments (me- lanophores) remain after preservation. The melano- phores on the yolk are found on the surface of the sac. On the embryo they initially appear on the dorsal part of the body and later on the head and tail. 3.2 Preadult phase 3.21 Embryonic phase See section 3.17. 3.22 Larvae and adolescent phase The size of a newly hatched larva of S. chiliensis is given by Barnhart (1927) as 3.75 mm. Newly hatched lar- vae of S. orientalis measured 4.1-4.3 mm (Harada et al. 1974). As a typical example of the development of the larvae of bonitos, a description of the larval and post- larval development of S. sarda is given below. The description is taken from a translation of Padoa (1956). Embryonic development is rapid and the larva hatches only a day after the closure of the blastopore. Figure 8. — The fertilized eggs of Sarda sarda at various developmental stages. (Note: The developmental stages were not defined by the authors.) (From Demir and Demir 1961.) 21 The mouth of the larva is not yet open, and the eye without pigment at hatching (Fig. 9a). The yolk sac and fin folds are large, and several oil droplets are present in the posterior part of the yolk sac. The anal opening is lo- cated a little less than midway between the head and tail. The pectorals are barely outlined. There is a total of about 50 body segments, including 15 preanal and 38 postanal segments. The yolk is reduced to less than half in a 4.6 mm speci- men 2 days after hatching (Fig. 9b). The mouth is open, the anus has moved forward, and the number of preanal segments is decreased from 15 to 11 and the postanal increased to 39. The pectorals are well developed and membranous; the hypurals are absent. Melanophores on the trunk are limited to the caudal ventral margin; other melanophores are present in the curve of the peritoneal cavity and a few are found on the profile of the snout. The yolk is nearly used up in a 4.68 mm specimen 4 days after hatching (not illustrated). The snout is con- siderably longer and there are well-defined teeth. There is a posteriorly directed triangular spine at the pre- opercular margin. The anus is displaced even farther forward to about the 9th segment. The pectorals are un- changed and the hypurals are still absent. The melano- phores persist on the caudal ventral margin; the number and size of melanophores are increased on the dorsal peritoneal walls and the head. The yolk is exhausted in a 4.20 mm specimen 6 days after hatching (Fig. 9c). The snout is a little more dis- tinctly pointed and is changing to the subconical form. The preopercular spine is pointed and evident; the pre- anal distance is about 34% of the total length. The pec- torals are membranous and the rounded margin reaches about the level of the anus; the hypurals are still absent. There are a few irregularly spaced melanophores on the ventral margin of the trunk. The melanophores persist on the peritoneal curve, along the thoracic girdle, and on the upper profile of the head; they are also present at the apex of the snout and in the middle of the mandi- ble. Melanophores are present on the apex of the snout, on the profile of the head, on the peritoneal curve, and along the thoracic girdle in a 7.2 mm specimen (Fig. 10c). Black pigments are almost completely lacking on the caudal trunk and limited to a few dispersed elements along the ventral margin. Two series of preopercular spines (an anterior of three and a posterior of six), of which the two central ones are very long, are now de- veloped; two other spines are present on the posterior margin of the otocysts. About 10 pointed teeth are on each side of the maxillary and the mandible. Outlines of about 13 rays, the future second dorsal and anal fins, can be counted. The preanal distance now represents 42% of the total length and the distance from the snout to the Figure 9. — Prelarvae of Sarda sarda: (a) 4.32-mm prelarva (from Padoa 1956, fig. 309); (b) 4.6-mm prelarva (from Padoa 1956, fig. 310); (c) 4.20-mm prelarva (from Padoa 1956, fig. 311). 22 2 n ^ 6 CM g ^.'■ , ? u> 3 E c a £ W to *C "? E H 3 fa W 23 insertion of the anal fin, 65Tr; there are about 20 seg- ments in the postanal region. In a 26.5 mm specimen, the preopercular spines are well-evident but small (Fig. lOf). The caudal fin is now forked; the pectoral and ventral fins are relatively small. The membrane that is continuous with the second dorsal and anal fins still unites the finlets. The first dorsal is covered with black pigments except for a clear space at the base of the 10th to the 13th rays; the other fins are transparent. Fine black pigments are on the snout, the nape, and the dorsal region of the trunk where six verti- cal bands, which extend from the dorsal edge to about the middle of the flanks, begin to appear. A dark hori- zontal line separates the pigmented dorsal portion from the ventral, unpigmented portion. In a 32 mm specimen the preopercular spines are smaller but still evident (Fig. lOg). The membrane still joins the finlets and the fin coloration is unchanged. The pigmentation on the head is increased and the six vertical bands are quite pro- nounced. Illustrations of postlarval S. sarda given by Vodianit- skii and Kazanova (1954) are also shown in Figure 10. Pinkas (1961) and Klawe (1961b) gave descriptions of postlarval S. chiliensis, and Jones (1960), Gorbunova (1963), and Harada et al. (1974) described larval and postlarval S. orientalis (Figs. 11, 12). Except for S. chiliensis and S. orientalis in the eastern Pacific, where there may be some overlap in the distri- bution of these two species, the problem of identifying and separating the larval and juvenile species within the genus does not occur because the species are allopatric. As for the larval forms of S. chiliensis and S. orientalis in the eastern Pacific, Klawe (1961b) stated that it would be impossible to separate the two species without a com- plete developmental series of both species. Klawe fur- ther noted that it should be possible to distinguish ju- venile S. chiliensis and S. orientalis from each other based on gill raker counts even at a relatively small size. Pinkas (1961) pointed out other characters (number of teeth on lower jaw and presence or absence of posterior gill teeth) that could be used to separate S. chiliensis >24 mm from similar sized S. velox ( = S. orientalis). Once the juveniles acquire the full adult complement of certain characters, which seems to be at a relatively small size (Pinkas 1961; Klawe 1961b), the summary of distinguishing characters (Table 1) for the four species of Sarda compiled by Collette and Chao (1975) indicates that the juveniles should be separable without difficulty. Sarda australis, S. chiliensis, and S, orientalis can be separated from each other on the basis of gill raker counts while S. sarda can be distinguished from all the others by the number of vertebrae. 3.3 Adult phase 3.31 Longevity See section 4.13. 3.32 Hardiness See section 3.52. 3.33 Competitors Sarda chiliensis The relationships of S. c. lineolata with the other species occupying the same areas off southern California are not clear. However, recent changes in the species composition of pelagic fish stocks off California shores may involve the bonito as well as Pacific sardine, Sar- dinops sagax, and northern anchovies, Engraulis mordax (Frey 1971). Presumably these changes would involve the ability of the various species to compete with one another for food and space. Behavior studies on S. c. lineolata indicate that they can compete very well with other species in obtaining food. In a fish tank containing Pa- cific barracuda, Sphyraena argentea; yellowtail, Seriola dorsalis; and tarpon, Megalops atlantica, S. c. lineolata was the first species to reach and ingest food tossed onto the surface (Magnuson and Prescott 1966). Magnuson and Prescott noted that the high speed of S. c. lineolata gives it an advantage over the other species in a "scramble" type of competition. Sarda orientalis In the Indian Ocean adult S. orientalis are often caught together with Euthynnus affinis, Auxis spp., Thunnus tonggol, and Scomberomorus spp. and pre- sumably compete with these species for food (Silas 1963). Demir (1963) stated that all predatory fishes and dol- phins are potential competitors of S. sarda, including Delphinus delphis ponticus, Tursiops truncatus, Scomber scombrus, Trachurus mediterraneus, Poma- tomus saltatrix in the Black Sea, and additionally, Scomber japonicus, Auxis rochei, and Euthynnus alletteratus in the Sea of Marmara. 3.35 Parasites, diseases, injuries, and ab- normalities Records of parasitism on bonitos include a trematode, Didymozoon pelamyzis, from the gill lamella of S. sarda from the Black Sea, Sea of Marmara, and the Medi- terranean (Dawes 1946; Demir 1963). Larvae of the cestode Callitetrarhynchus gracilis have been found in the body cavity and the isopod Livoneca sp. from the gill lamellae of S. sarda (Postel 1954). Parasitic copepods Caligus sarda (Pearse 1952) and Ceratocolax euthynni (Vervoort 1971) have been found on S. sarda. Silas (1967) and Silas and Ummerkutty (1967) produced a detailed listing of the parasites of scombroid fishes, including Sarda (Table 13). Two stripeless S. c. lineolata have been recorded from waters off La Jolla, Calif. (Matsumoto et al. 1969). These specimens possessed all the meristic characters typical of S. c. lineolata except that they lacked stripes. 24 Figure 11.— Postlarval Sarda chiliensis: (a) 16.7 mm fork length (from Pinkas 1961, fig. 4); (b) 33.0 mm fork length (from Pinkas 1961, fig. 5); (c) 42 mm (from Klawe 1961b, fig. 3); (d) 160 mm (from Klawe 1961b, fig. 4). 25 E ~ lis »j _M .3 "p. to . O •* ft ov £ ~ E * s a C ■o -' cd — SB a . _ St E c 2? S — > *a a « — a. E _- E s - a _• -» a e a ; c £ ■ » 3 si "C - c o -a . Ill o a * "? S t -a "5 a « e to S g 7 a E o 31 a — Zx3 26 Table 13.— Parasites of Sarda. (Modified from Silas 1967; Silas and Ummerkutty 1%7.) Monogenetic Digenetic Copepods trematodes trematodes Cestodes Sarda chiliensis Caligus bonito Caligus mutabilis Pseudocycnus appendiculatus Sarda orientalis Caligus bonito Capsala ca ballerio Bucephalopsis cybii Caligus coryphaenae Caligus productus Sarda sarda Caligus bonito Capsala pel imydis Aponurus tschugunowi Grillotia erinaceus Caligus mutabilis Hexostoma pricei Atalostrophion sardae Lacistorhynchus Caligus pelamydis Hexostoma thynni Bucephalopsis arcuata tenus Caligus productus Dinurus barbatus Hirudinella clavata Lecithochirum caudiporum Lecithochirum texanum Nematobothrium pelamydis Opecoelides vitellosus Rhipidocotyle angusticollis Tormopsolus orientalis? Unitubulotestes sardae Scolex pleuronectis Tentacularia bicolor Tentacularia coryphaena Tetrarhynchus megabothrium Tetrarhynchus scomber-pelamys Tetrarhynchus sp. '' Species incertae sedis. "Indicates doubtful record. Forms unidentifiable. 3.4 Nutrition and growth 3.41 Feeding Magnuson and Heitz (1971) examined the relation be- tween the gill raker morphology and the food habits of scombrid fishes, including S. c. lineolata. In the past, the high diversity of food organisms in the stomachs of scom- brids gave rise to the opinion that scombrids are non- selective feeders. However, Magnuson and Heitz pointed out that selectivity does exist in terms of food size in scombrid food habits in that the stomach contents of small and large fish of the same species were dissimilar. They suggest that larger predators have a reduced ability to catch small prey (crustaceans) because of a relatively large gap between the gill rakers. Among scombrids of the same size S. c. lineolata and S. orientalis had the largest gill gaps (1.8-3.3 mm). The relative unimpor- tance of crustaceans in the food habits of S. c. lineolata noted by Pinkas et al. (1971) could be related in part to the large gill gaps in this species. The data presented by Magnuson and Heitz (1971) also indicate a relationship between the number of gill rakers and the gill raker gap, i.e., the greater the number of gill rakers, the smaller the gill raker gap. Sarda orientalis have fewer gill rakers than S. chiliensis (Collette and Chao 1975), and as Magnuson and Heitz's data show, the gill raker gap is greater in S. orientalis. Therefore, it would be expected that the diet of S. c. lineolata would contain a larger proportion of smaller organisms than that of S. orientalis. 3.42 Food Sarda australis Only a few miscellaneous observations are available on the feeding habits of the Australian bonito. Particularly during the winter months, S. australis occurs commonly in schools in the inshore coastal waters of Queensland and feeds on hardyheads, Pranesus ogilbyi; pilchards, Sardinops neopilchardus; and anchovies, Engraulis australis (Grant 1972). Munro (1958) indicated that they also feed on mackerel scad, Trachurus mccullochi. Sarda chiliensis A detailed study of the food habits of S. c. lineolata was conducted by Pinkas et al. (1971) based on a total of 1,498 stomachs collected in 1968 and 1969 from fish cap- tured in nearshore waters of southern California and Baja California. Their study clearly showed that the northern anchovy, Engraulis mordax, was the major food item in the diet of S. c. lineolata. The common squid, Loligo opalescens, ranked next in importance, and mis- cellaneous fishes and a few crustaceans made up the small remainder of the diet of S. c. lineolata (Table 14). 27 Table 14.— Food of Sarda chiliensis lineolata, 1968 and 1969. (From Pinkas et al. 1971.) Frequency Percent Percent Volume Percent of frequency of Food items Number number (ml) volume occurrence occurrence Fishes Engraulidae Engraulis mordax 4,159 75.5 11,356.4 75.9 462 56.3 Scomberesocidae Cololabis saira 27 0.5 190.8 1.3 16 1.9 Gadidae Merluccius productus 24 0.4 6.8 <0.1 15 1.8 Carangidae Trachurus symmetricus 16 0.3 185.6 1.2 13 1.6 Sciaenidae Genyonemus lineatus 1 <0.1 38.0 0.3 1 0.1 + Seriphus politus 3 +0.1 1.3 <0.1 2 0.2 + Embiotocidae Brachyistius frenatus 1 <0.1 10.0 <0.1 1 0.1 + Cymatogaster aggregata 1 <0.1 14.0 0.1 1 0.1 + Zalembius rosaceus 1 <0.1 9.0 <0.1 1 0.1 + Scorpaenidae Sebastodes spp. 12 0.2 5.8 <0.1 5 0.6 Stromateidae Peprilus simillimus 7 0.1 + 195.0 1.3 6 0.7 Bothidae Citharichthys sordidus 1 <0.1 0.6 <0.1 1 0.1 + Unidentified fishes 765 13.9 223.8 1.5 185 22.5 Subtotal, fishes other than Engraulidae 859 15.6 880.7 5.9 247 30.1 Subtotal, all fishes 5,018 91.1 12,237.1 81.8 709 86.4 Cephalopods Loligo opalescens 448 8.1 2,690.7 18.0 207 25.1 Onychoteuthis boreali-japonicus 1 <0.1 3.7 s 9.9 I 1 0.1 + Unidentified cephalopods 17 0.3 + 16 2.0 Crustaceans . 0-2 Pleuroncodes planipes 4 <0.1 5.6 4 0.5 Crab megalops larvae 3 <0.1 0.1 1 0.1 + Unidentified animals 19 0.3 + 4.7 I 17 2.0 Subtotal, all invertebrates (except Loligo opalescens) and unidentified animals 44 0.8 24.0 0.2 39 4.8 Totals 5,510 100.0 14,951.8 100.0 — — There was a seasonal variation in the importance of the common squid in the diet of S. c. lineolata. Squid was most important in the first and second quarters of the year but was less important in the third quarter of the year. This seasonal pattern of the importance of squid in the bonito's diet was related to the seasonal con- centrating behavior of squid during its reproductive cy- cle. Sarda orientalis Information on the food and feeding habits of S. ori- entalis in Japanese waters is fragmentary. Yabe et al. (1953) examined 18 stomachs from juvenile S. orientalis in southern Kyushu waters. Fifteen of the 18 stomachs were empty, 2 of the stomachs contained Engraulis japonicus remains, and 1 stomach contained unidentifi- able fish remains. Yokota et al. (1961) examined stomachs of 24 S. orientalis caught in about the same area and found a total of five saury, one Sphyraena sp., one squid, and one carangid in the stomachs. Kumaran (1964) investigated the food habits of S. ori- entalis in the Indian Ocean based on 43 specimens rang- ing in length from 85 to 305 mm. The most important single food item was Anchouiella commersonii. Fishes of lesser importance in the diet were Leiognathus insidi- ator, Decapterus russelli, and Sardinella sp. Kumaran noted that the variety of food organisms was smaller in the stomachs of S. orientalis as compared with Euthyn- nus af finis and Auxis. He gave as the reason for this the fact that all the S. orientalis specimens were collected from a single locality. Sivasubramaniam (1969) looked at the food-bearing stomachs of 11 S. orientalis caught in the waters around Sri Lanka. Nearly 60% by volume of the food consisted of a clupeoid fish, 18% cephalopod mollusk (squid and octopus), 15% decapod Crustacea, and 1% miscellaneous items including unidentified fishes. Sarda sarda More information is available on the food and feeding habits of S. sarda, including information on larval and 28 juvenile stages. Larval S. sarda about 5 mm long start active feeding even before the yolk sac is completely ab- sorbed. Larvae >6-7 mm feed on relatively large organ- isms, including fish larvae, but prefer copepods. Ju- veniles >18-20 mm feed on the larvae of anchovy, horse mackerel, and S. sarda (Mayorova and Tkacheva 1959). Zusser (1954) reportea a Pseudocalanus in the stomach of a 7.2 cm juvenile and a 4.5 cm long anchovy in a 8.7 cm juvenile. Demir (1963) found a single 3.5 cm horse mackerel from the stomachs of 44 juvenile S. sarda rang- ing in length from 6.5 to 16 cm taken in the Black Sea, the Sea of Marmara, and the Bosporus from 1957 to 1959. Because of the preponderance of empty stomachs, Demir believes that the juveniles, like the adults, regurgitate their food at the moment of capture. Demir (1963) stated that adult S. sarda is an insati- able predator that feeds diurnally. Feeding in Turkish waters is much more vigorous in the early morning and towards evening. Demir noted that the feeding season is usually from the second half of April to the end of Octo- ber in Turkish waters. Sarda sarda adults primarily feed on smaller schooling fishes, the species depending on the locality (Table 15). In the western Atlantic in the Gulf of Maine, S. sarda prey on mackerel, alewives, menhaden, other small fishes such as launce and silverside, and squid (Bigelow and Schroeder 1953). Boschung (1966) examined the stomachs of 52 S. sarda taken during a fishing tourna- ment in the Gulf of Mexico and noted that the fish had fed on a variety of fishes and invertebrates. They in- Table 15. — Food of Sarda sarda in the Black Sea, Sea of Marmara, and the eastern Atlantic. (From Berg et al. 1949; Postel 1954; Zusser 1954; Slastenenko 1956; Ionescu et al. 1958; Demir 1963.). Black Sea Engraulis encrasicholus Spratella sprattus phalerica Ammodytes cicerellus Scorn ber scorn brus Trachurus mediterraneus Mullus barbatus Pomatomus saltatrix (young) Sarda sarda (young) Mugilspp. Atherina spp. Gobiidae Sea of Marmara Important food items Engraulis encrasicholus Spratella sprattus phalerica Scorn ber scombrus Scomber colias Less important food items Smarts alceda. Trachurus trachurus Clupea pilchardus Ammodytes cicerellus Trachurus mediterraneus Mullus barbatus Atherina spp. Boops boops Atlantic Ocean near Dakar, Senegal Sardinella sp. Engraulis sp. Scomber colias Ammodytes sp. Planktonic crustaceans Caprella Euphausiidspp. eluded: a clupeid (probably Harengula pensacolae) harvestfish, Peprilus paru; spot, Leiostomus xanthurus anchovies, Anchoa sp.; mackerel, Scomberomorus sp. sea robin, Prionotus sp.; squid, Loligo sp.; shrimp, Penaeus sp.; and unidentified fishes. Sarda sarda can swallow relatively large prey and the adults and juveniles are both cannibalistic. There is a record of a 38 cm S. sarda from the stomach of a 63 cm fish (Zusser 1954). 3.43 Growth rate Sarda australis There is almost nothing in the literature on the age and growth of S. australis except that they grow to 91.4 cm (36 in) (Ogilby 1954). Sarda chiliensis In the waters between southern California and north- ern Baja California S. c. lineolata spawns between Janu- ary and May and the pelagic fertilized eggs take about 3 days to hatch. Young S. c. lineolata are first observed in the early summer when they are 15.7-25.4 cm (6-10 in) and by the following spring they are about 38.1 cm (15 in) long and weigh from 0.7 to 0.9 kg (1.5 to 2 lb). In the fall these fish may weigh 1.4 kg (3 lb) or more and by the fol- lowing year in May, they may weigh 2.7-3.2 kg (6-7 lb). When they are 4 yr old, they are 71.1 cm (28 in) and weigh about 5.4 kg (12 lb) (Frey 1971). These data are probably the result of random observations and not a systematic study. Kuo (1970) made a detailed study of the growth of S. chiliensis from southern California waters and from Peru based on otoliths (Fig. 13). Kuo fitted the von Bertalanf- fy growth function to his data and found that the length- at-age of bonito from the two areas did not differ for age 1, 2, and 6 fish but that there was a significant difference in size offish 3-5 yr old (Table 16). The growth equations obtained for the northern and southern populations, re- spectively, were: L t = 2,661 [1 - e -0.038lt+ 06011 and L t = 1,014 [1 -e- 0154 '^ where L, is fork length in millimeters and t is actual age in years. Campbell and Collins (1975) also determined the age and growth of the northern population of S. chiliensis based on a sample of 3,139 fish ranging in length from 23 to 79 cm landed at the canneries in San Pedro, Calif. These fish were caught in an area between central Baja California and Point Conception, Calif., from 1968 to 1974. Like Kuo (1970), Campbell and Collins used otoliths to age the fish and used the von Bertalanffy growth function to describe the growth as shown below: 29 90 "0 60 x 50 I- © 20 CAMPBELL 8 COLLINS (1975) /, X Sarda chiliensis lineolato . /Y \e m ur k, i hKUO 1970) .' . Sarda chiliensis chiliensis 5 AGE Figure 13. — Growth of Sarda chiliensis from southern California and Peru. (Note: The abscissa of the Campbell and Collins curve is age class.) (From Kuo 1970; Campbell and Collins 1975, fig. 4.) Table 16. — Age-length data for the subspecies of Sarda chili msi's. (From Kuo 1970, table 14.) Mean Confidence length Standard interval Age Species (mm) error (mm) N 1 lineolata 152 0.95 150-154 222 chiliensis 151 1.99 147-155 141 2 lineolata 252 1.28 249-254 219 chiliensis 256 2.76 250-261 137 3 lineolata 347 1.70 344-351 212 chiliensis 364 6.02 352-376 43 4 lineolata 433 2.26 429-438 182 chiliensis 480 7.01 466-494 26 5 lineolata 509 3.19 503-515 133 chiliensis 550 7.27 535-565 25 6 lineolata 585 4.79 576-595 74 chiliensis 604 9.18 585-623 24 7 lineolata 652 13.00 625-678 35 chiliensis — — — 8 lineolata 756 12.10 728-783 9 chiliensis — — — 'lineolata = Sarda chiliensis lineolata. [Northern Hemisphere] chiliensis = Sarda chiliensis chiliensis. [Southern Hemisphere] L, = 76.87 [1 - e •0.6215it-H0.410>' where L t is fork length in centimeters and t is age at cap- ture, which is about 0.375 yr (September) older than the given age. The length-age relationship predicted by the curve for fish taken in the fishery is shown below: ge Fork length (cm) 1 44.9 2 59.7 67.6 71.9 74.2 75.4 The growth curve is also shown in Figure 13. The maximum size of S. chiliensis in the Northern Hemisphere is about 101.6 cm (40 in) and about 11.3 kg (25 lb). There is an unverified record of a 16.8 kg (37 lb) specimen, but bonito over 11.3 kg are rare (Frey 1971). In light of these observations, the L r of 2,661 mm obtained by Kuo (1970) for the Northern Hemisphere bonito seems unreasonably large. On the other hand, as noted by Campbell and Collins (1975), the L„ predicted by the von Bertalanffy growth function as fitted to their data was less than what they had observed in the California fish- ery and much less than the maximum length of S. chiliensis (101.6 cm) in the literature. They noted that the discrepancy could have been caused by the lack of fish larger than 79 cm in their sample. It is of interest also that the growth curves obtained by Kuo (1970) and Campbell and Collins (1975) for the northern population of S. chiliensis are quite different. The results obtained by Campbell and Collins indicate a rapid growth rate during the first 3 yr and a tapering off to a relatively low rate thereafter. Kuo's growth curve on the other hand shows only a slight curvature and sug- gests almost linear growth for the northern bonito population (Fig. 13). The age-length data given by Campbell and Collins (1975) and Kuo (1970) are also quite different. The lengths at the different ages as given by Campbell and Collins are larger for all the ages than those given by Kuo. The maximum size of 5. chiliensis in the Southern Hemisphere is not well defined. Barrett (1971) sampled the fish landed in the Chilean fishery and noted that the maximum size of fish landed was 74 cm. In an earlier study Buen (1958) found fish as large as 79 cm. Mann (1954) stated that Chilean bonito grows larger than 80 cm. Sarda orientalis Very little information is available on the age and growth of S. orientalis. Yabe et al. (1953) presented data on the mean lengths of six samples of S. orientalis rang- ing in length from 161 to 348 mm that were sampled at Aburatsu, Japan, over an irregular period from 26 August to 17 October 1950 (Table 17). The mean lengths offish from the six samples were plotted against sampling date, and a straight line was drawn by eye through the data points (Fig. 14). If the line is a reasonable representation of the growth of S. orientalis between 161 and 348 mm, it indicates a mean growth of about 2.6 mm/day. Harada et al. (1974) made observations on the growth of larval and juvenile S. orientalis in artificial fertilization experi- ments. They determined that the fertilized eggs hatched in about 50 h in water temperature ranging from 20° to 24°C and that the newly hatched larvae were 4.1-4.3 mm in total length. The larvae grew to 14 mm in total length 30 Table 17. — Lengths of Sarda orientalis landed at Aburatsu, Japan in 1950. (From Yabe et al. 1953, table 11.) Body length (mm) Table 18.— Monthly length data for Sarda sarda from the Black Sea. (Data from Demir 1963.) (Note: The author did not define the length dimension.) Date 1NO. 01 fish Range Mean Author July Aug. Sept. Oct. 17 205-221 214 r h(cm) - - Aug. 26 S Sept. 1 2 241-249 245 Zusser(1954) — 19-27 27-29.5 25-36 Sept. 2 3 233-243 239 Tkacheva (1958) — 21-33 27-36 36-40 Sept. 8 5 161-270 239 Mayorova and Tkacheva (1959) — 21-33 27-37 36-41 Sept. 9 10 254-277 263 Demir (1963) 6 L6 12-32 25-38 33-42 Oct. 17 5 333-348 339 AUG. Figure 14.- SEPT. OCT. -Growth of juvenile Sarda orientalis in Japanese waters. (Data from Yabe et al. 1953.) in 10 days, to 74 mm in 20 days, to 106 mm in 30 days, to 219 mm in 42 days, and to 290 mm in 99 days after hatching. This growth rate represented the fastest growth under the conditions provided in the experi- ment. The growth rate of larval and juvenile S. ori- entalis indicated by these experiments was about 2.9 mm/day during the 99-day period and compares favor- ably with that suggested by the data presented by Yabe et al. (1953). Harada et al. (1973) presented some data on the growth of S. orientalis kept in artificial enclosures. Thirty-three S. orientalis 40 cm long and 675 g in weight grew to 50 cm and 1,500 g in about 4 mo. Regarding the maximum size of S. orientalis in the In- dian Ocean, the large specimens off the southwest coast of India are generally less than 70 mm long (Silas 1964). Smith (1949) stated, however, that S. ehihensis (= S. orientalis) grows to at least 101.6 cm (40 in). In Japanese waters S. orientalis grows to about 80 cm and 1.5-3.0 kg (Kishinouye 1923). Sarda sarda Various investigators have presented monthly length data for young S. sarda from the Black Sea (Table 18). The data were given as the ranges of lengths sampled during monthly periods, and the midpoints of the length ranges were plotted to represent the growth of 3-42 cm S. sarda in the Black Sea (Fig. 15). As would be expected, there is some variation in the data presented by the various investigators, but the variation was relatively small. In gross terms these data suggest that S. sarda grew about 252 mm in 90 days (7.5-11 cm in July to 30.5- 38.5 cm in October) or at a growth rate of 2.8 cm/day. Demir (1963) summarized the age-length relation of S. sarda determined by various authors (Table 19). Subse- quent to the publication of Demir's paper, Kutaygil (1967) published some data on age-length determina- tions on S. sarda (Table 20). Kutaygil used otoliths to age the fish and also back-calculated the lengths of the fish at various ages. He noted that bonito from different year classes had different growth rates. Zusser (1954), in contrast to the other investigators, gave age-length relations for fish up to 9 yr of age. Demir (1963), however, remarked that the age-length relations given by Zusser were probably erroneous. Although sev- eral age-length relations for S. sarda are available, as far as it is known, no one has tried to fit a growth curve to these data. 30 25 e o J 20 UJ x x ZUSSER0954) Q TKACHEVA (1958) v 7 MAYOROVA a TKACHEVA0959) o o DEMIR (1963) a A MEYER (1956) JULY AUG SEPT OCT. Figure 15. — Growth of juvenile Sarda sarda in the Black Sea. :si Table 19.— Age-length relation in Sarda sarda. (From Demir 1963, table 4.) (Note: The length dimension was not defined by the author.) Age Author 1 2 3 4 5 6 7 8 9 - Length (cm) - Zusser (1954) 25-37 33-50 42-54 50-62 56-67 60-70 63-76 70-78 74-85 31.5 41.5 48.8 56.2 61.2 66.5 69.5 73.5 76.2 Numann (1955) 38-41 53-57 60-64 — — — — — — Nikolskii (1957) 35.3 55.1 64.2 72.5 — — — — — Tiirgan (1958) 30-40 50-55 55-60 60-65 — — — — — Nikolov (1960) 38.85 52.6 60 67 (74-75) — — — — Table 20. — Calculated lengths for each age of Sarda sarda. (From Kutaygil 1967, table 26.) (Note: The author did not define the length dimension.) No. of fish Age Sample 1 2 3 4 1957 II (1955) 7 44.0 59.1 — in (1954) 22 45.4 57.9 64.8 — IV (1953) 4 47.6 59.2 64.5 68.7 Granc mean = 45.1 58.3 64.9 N = 33 33 26 Jan. 1958 IV (1954) 5 42.9 53.3 60.6 64.2 Feb. 1958 IV (1954) 18 41.1 52.9 59.2 63.8 The largest S. sarda is about 85 cm in the Black Sea. Of S. sarda in the western Atlantic, Bigelow and Schroeder (1953) stated that they grow to about 91.4 cm (36 in) and 4.5-5.4 kg (10-12 lb). Hammond and Cupka (1975) noted that S. sarda seldom exceeds 6.8 kg (15 lb). 3.5 Behavior 3.51 Migrations and local movements See also section 5.3. The seasonal migrations of S. sarda from the Aegean Sea through the Sea of Marmara to the Black Sea have been well documented. The migration routes were pre- sumably determined by the development of the fisheries in these areas and were verified by tagging experiments (Demir 1963). The migration from the Aegean Sea to the Black Sea starts toward the end of April and lasts to the beginning of June or later. The return migration from the Black Sea to the Aegean Sea usually starts in Septem- ber and lasts to the end of November. Some of the bonito schools returning from the Black Sea may stay in the Sea of Marmara but others continue on to the Aegean Sea. The S. sarda in the Adriatic Sea apparently make a north-south migration within that sea down to around the Greek islands (Belloc 1954). The bonitos leave the upper Adriatic in November-December and travel along both shores of the Adriatic, arriving in Greek waters in February-March. The return migration starts around August. In addition, there apparently are schools of bonito that remain the whole year round in Greek waters and in waters along the east coast of Sicily. The relation of the bonitos in the Aegean Sea-Sea of Marmara-Black Sea complex with the Adriatic Sea bonito and with the bonitos in the other parts of the Mediterranean is not clear. Turkish investigators imply that the stock in the Aegean Sea-Sea of Marmara-Black Sea complex originates in the Aegean Sea and the Sea of Marmara (Demir 1957) and remain within this complex. In the eastern Pacific, over 11,200 S. c. lineolata have been tagged and released since 1968 along the coast from Monterey Bay, Calif., to Cape San Lazaro, Baja Cali- fornia (Collins and MacCall 1977). Recoveries of tagged fish have indicated that they move randomly in local California waters, although there was a definite move- ment down the southern California coast in the winter and a northward movement in the late summer and fall, apparently in response to changes in sea temperature. Their tagging study also indicated that heated water dis- charges from coastal electric generating stations strongly influence the migration of young S. c. lineolata. Fish tag- ged in heated water plumes either remained in the area of the discharge or tended to migrate to another heated discharge area. 3.52 Schooling All species of Sarda generally appear to be schooling fish. Sarda australis commonly occur in schools in in- shore coastal waters of Queensland, Australia (Grant 1972). Frey (1971) stated that the S. chiliensis of the Northern Hemisphere is a pelagic schooling fish. Tominaga (1943), however, observed that S. orientalis in Japanese waters do not aggregate densely, rarely come to the surface of the sea, and always swim around reefs or near a cape where the current is strong, and that they never go out to the high seas. Conflicting observations are available on schooling of S. orientalis in the Indian Ocean. Silas (1963) stated that schools of adults and young appear along the southwest coast of India. Si- vasubramaniam (1969), however, noted that S. ori- entalis are very seldom seen in surface schools of mixed tunas and when they are caught from mixed schools, the catch never exceeds six fish. All of the observations on S. sarda indicate it is a schooling fish. Demir (1963) stated that S. sarda gathers in dense schools of many thousands of fish of about the same size. Sarda sarda in the western Atlantic is a schooling fish, traveling in large aggrega- tions. It is usually found at the surface, although oc- casionally it is caught near the bottom (Idyll and de 32 Sylva 1963) . Of 5. sarda that occurs in the Gulf of Maine, Bigelow and Schroeder (1953) wrote, "The bonito is a strong, swift, predaceous inhabitant of the open sea and like its tribe travels in schools." 3.53 Responses to stimuli A detailed, systematic study on the behavior of captive S. c. lineolata in a large fish tank at Marineland of the Pacific, Palos Verdes, Calif., was made by Magnuson and Prescott (1966). The courtship and spawning be- havior of S. c. lineolata, as reported by Magnuson and Prescott, was discussed earlier in sections 3.11 and 3.13. Further results of their observations are detailed below. About 25 S. c. lineolata out of 60 that were caught near Marineland of the Pacific became established in the tank and were used in the observational program. Of the 25 survivors, 10 fish lived for 38 mo in captivity. The tank used was part of a public display and contained more than 800 other fishes of over 40 species. Part of the daily routine included five 15-min shows in which a diver entered the tank to feed the fish. Magnuson and Pres- cott noted that the S. c. lineolata "appeared habituated to show announcements and background music as well as to sounds made by tapping on the tank windows and sides." These observations indicate that S. c. lineolata are able to adapt to various environmental conditions. In the Marineland of the Pacific tank, S. c. lineolata swam continuously against the current averaging 88.2 cm/s at a tail-beat frequency of 1.42 beats/s when not feeding or courting. Sarda c. lineolata apparently are less powerful swimmers than skipjack and yellowfin tunas. In fish about the same length (bonito, 57 cm; skipjack tuna, 57 cm; yellowfin tuna, 52 cm) Magnuson and Prescott noted that at four tail beats/s S. c. lineolata traveled only 170 cm/s whereas Katsuwonus pelamis averaged 230 cm/s and T. albacares averaged 240 cm/s. Magnuson and Prescott observed nine miscellaneous behavior patterns in S. c. lineolata: mouth closure (long), mouth closure (short), snap, yawn, quick swim, lean, bend, jerk, and defecation. They discussed in detail the possible functions of all these miscellaneous behavior patterns. They hypothesized that mouth closure (long) movements could have been associated with olfaction or gill ventilation; snaps following a yawn or a quick swim with a drinking movement; leans may have a function as a social releaser in schooling; and bends and jerks may be associated with food passing through the alimentary canal. No such detailed behavioral observations have been Table 21.— Sex ratio of Sarda sarda landed in Spain. (From Rodriguez-Roda 1966.) Ratio Year Month Port Males Females (Male:Female) 1963 June Tarifa 17 18 1:1.06 1964 May Barbate 57 20 1:0.35 1964 June Barbate 25 24 1:0.96 1964 July Barbate 18 24 1:1.33 made on other species of Sarda. Inoue et al. (1967) were able to maintain S. orientalis up to 438 h in a pool. Inoue et al. (1970) determined that S. orientalis was negatively phototactic to both sunlight and artificial light. The swimming speeds of the fish (size not given) ranged from 0.3 to 0.56 m/s. 4 POPULATION 4.1 Structure 4.11 Sex ratio Sarda australis No information. Sarda chiliensis Kuo (1970) obtained sex ratio data on S. c. lineolata that were caught from San Diego waters during a 1-yr pe- riod between 1964 and 1968. The percentage of females in the monthly samples varied from 37.5 to 70.3% and aver- aged 49.9% for the year. He found that the monthly sex ratios did not differ significantly from 1:1 except for the sample from the month of May. The percentage of female S. c. chiliensis from eight samples of 100 bonito each obtained from the commer- cial catch landed at Iquique, Chile, varied from 47 to 65% between September 1968 and October 1969 (Barrett 1971). Barrett noted that more females were present in the catch during September-October spawning season, but stated that more data were needed to verify this observation. Sarda orientalis No information. Sarda sarda Rodriguez-Roda (1966) determined the sex ratio of four samples of S. sarda landed at the ports of Barbate and Tarifa, Spain (Table 21), in 1963 and 1964. The females in the samples ranged from 26 to 57.1%. Postel (1955b) presented data on the monthly sex ratio of S. sarda from the eastern tropical Atlantic (Table 22). In the Gulf of Mexico a sample of 52 S. sarda was composed of 31 males and 21 females (Boschung 1966). 4.12 Age composition See section 4.13. 4.13 Size composition Sarda australis There is no information on the size or age of the fish ex- cept for generalized observations. The species grows to 33 Table 22. — Sex ratio of Sarda sarda from the eastern tropical Atlantic. (From Postel 1955b.) Ratio Month Males Females (Male:Female) January 13 12 1:0.42 February 38 30 1:0.79 March 65 70 1:1.08 April 142 115 1:0.81 Mas- 148 172 1:1.16 June 1 1 1:1 > o z July — — — August — — — ui September — — — o 111 October 3 8 1:2.67 u_ November 7 5 1:0.71 z December 5 6 1:1.20 111 o K Ul Q. Total 422 419 1:0.99 about 90 cm (3 ft) but the average size of fish caught is about 40 or 45 cm (16 or 18 in) (Marshall 1964). Grant (1972) stated that the usual size taken is 1.8-2.3 kg (4-5 lb). Sarda chiliensis The length-frequency distribution of S. c. lineolata caught by sports fishermen from southern California waters shown in Figure 16 was prepared from data pro- vided by C. M. Kuo 4 and is based on part of the S. c. lineolata sample he used in his study (Kuo 1970). Kuo measured 929 S. c. lineolata from southern California waters ranging in length from 331 to 750 mm. Both the male and female length-frequency distributions were similar: a single prominent mode was present between 510 and 540 mm. MacCall et al. (1976) presented S. c. lineolata length- frequency data for the California commercial and party- boat fisheries in 1973 (Fig. 17). They stated that various segments of the fishery exploit different parts of the bonito population, which indicates an uneven geo- graphical distribution of various age groups. It was indi- cated that, generally, the older fish were more available near Mexico and in offshore waters, although large fish were also taken in the fall off Santa Barbara. The party- boat fishery caught bonito <60 cm but the long-range party boats fishing off Mexico took older fish. The com- mercial fishery took larger fish. It is of interest that the modal size taken by the party boats during the period 1964-68 was larger than that taken in 1973. Recruitment strength is highly variable in southern California waters, and because the party-boat fishery takes only young fish, the catch per effort and the size composition of the catch reflect this variability and may account for the differ- ence in modal sizes (MacCall 3 ). Barrett (1971) presented bonito length-frequency data from Chilean landings and noted that the larger bonito, HI Presently with Oceanic Institute, Makapuu Point, Waimanalo 96795. MacCall, A. D., California Department of Fish and Game, c/o Na tional Marine Fisheries Service, La Jolla, CA 92038, pers. commun February 1978. FEMALES (N = 2I2) ^tH 45 j=tl 20 10- MALES (N = 202) r-TTh rrhJrfr- □d 35 40 45 50 55 60 65 LENGTH (cm) Figure lf>. — Length-frequency distribution of Sarda chiliensis lineolata captured in the sport fishery in southern California. U.S. COMMERCIAL CALIFORNIA A N= 1,502 ' W= 6 45 lbs i 1973 MEXICO N = 365 W = 7.70 lbs vIEAN LENGTHS OF AGE GROUPS I m nz zvrzn U.S. PARTYBOATS LOCAL l\ N = 1,235 I W= 2.87 lbs y ' LONG RANGE OFF MEXICO N = II6 W= 8.79 lbs 30 40 50 60 70 80 LENGTH (cm) Figure 17. — Length-frequency distribution of Sarda chiliensis lineolata taken by the California commercial party-boat fisheries. (From MacCall et al. 1976, fig. 9.) in about the 62 cm modal group, made up most of the catch in September-October 1968 (Fig. 18). Most of the bonito in the April, August, September, and October 1969 landings, however, were smaller, 48-52 cm, and the larger fish were present only in July and August. Barrett 34 56 60 64 LENGTH (cm) Figure 18. — Length-frequency distribution of Sarda chilienss chili- ensis sampled from the commercial landings at Iquique, Chile, September 1%H to October 1%!). Numerals in upper left corner of each panel indicate the month and year, number offish, and number of samples. Unshaded distribution is from Buen (19S8). (From Bar- rett 1971, fig. 4.) further noted that the preponderant 72-74 cm modal group in 1953 reported by Buen (1958) was almost absent in the 1968 and 1969 samples, which indicated that the older fish were no longer present in the fishery in 1968 and 1969. Vildoso (1962) and Barrett (1971) determined length- weight relationships for the southern S. chiliensis population and Campbell and Collins (1975) for the northern population (Table 23). In computing length- weight relationships, many investigators use a logarith- mic transformation to linearize the data. Beauchamp and Olson (1973) pointed out that a bias is inherent in this procedure because the largest values are compressed on the logarithmic scale and provided a procedure to cor- rect for this bias. As they pointed out, corrections for this bias have been outlined in the past but seldom used in practice and it is most likely that the length-weight re- lationships in Table 23 have not been corrected for bias. Sarda orientalis Sivasubramaniam (1969) presented length data for S. orientalis from the Sri Lanka fishery (Fig. 19). The aver- age size of adult S. orientalis taken off southwest India was about 45 cm (Silas 1963). Data on the size composition of the stock of S. ori- entalis in Japanese waters are almost nonexistent. Yabe -et al. (1953) presented measurement data on a small sample of S. orientalis landed at Aburatsu, Japan (Table 17). Sivasubramaniam (1966) determined the length- weight relationship for S. orientalis from the waters around Sri Lanka in the Indian Ocean (Table 23). The length-weight relationship is based on a sample of 25 fish, and although Sivasubramaniam did not give the size range of the fish, his figure 14 indicates that the fish ranged from about 24 to 51 cm. Sarda sarda Black Sea landings of S. sarda in the spring of 1955 were dominated by a group of fish centered at a length of 45 cm (Fig. 20). In the spring of the subsequent 2 yr, the same group of fish (1956, 55 cm; 1957, 60 cm; 1958, 65 cm) dominated the catch. This group of fish was the result of a strong year-class that originated in 1954 (May- orova and Tkacheva 1959). Artuz (1959) also noted the same phenomenon. He sampled the landings at the Istanbul fish market and found that 3-yr-old fish (1954 Table 2.1. — Predictive length-weight relationships, W = ai b , of Sarda. (Note: Various length and weight units were used in the original regressions, shown in the table. For ease in comparison, the constants were converted where needed so that the regressions are in terms of weight in grams and length in centimeters.) Species Sex Size range of fish No. of Weight Length fish (g) (cm) a b Source 595 40-73 0.0118 3.02 Barrett (1971) 25 — 0.0152 2.958 Sivasubramaniam (1966) 165 — 40-55.5 0.0149 2.971925 Rodnguez-Roda (1966) 0.009088 3.09749 Campbell and Collins (1975) 2,824 200-7,675 29-77 0.009611 3.08338 Campbell and Collins (1975) 0.009376 3.08962 Campbell and Collins (1975) 565 — — 0.006491 3.19 Vildoso (1962) 513 — — 0.006311 3.19 Vildoso (1962) S. c. chiliensis S. orientalis S. sarda S. c. lineolata S. c. lineolata S. c. lineolata S. c. chiliensis S. c. chiliensis M Sexes com- bined M F 35 WEST COAST N = 20 SOUTH COAST N = 27 18-22 22-26 26-30 30-34 34-38 38-42 42-46 46-50 50-54 FORK LENGTH (cm) Figure 19. — Length-frequency distribution of Sarda orientalis around Sri Lanka (Ceylon) (1967-68). (From Sivasubramaniam 1969, fig. I.) O 40 60 I 1 1 I 1 1 1955 N = 1,286 A II h . 1 - 1956 N = 1,149 - 1957 N = 10,881 ! i i i r - 1958 N = 5,167 a : 1 I : _ \ ■ 30 35 40 45 50 55 60 65 70 75 LENGTH (cm) Figure 20. — Length-frequency distribution of Sarda sarda in the Black Sea, spring 1955-58. (Data from Mayorova and Tkacheva 1959, table 1.) year class) dominated the landings in May 1957. His sample showed that the May 1957 landings were com- posed of 10.5% age I fish (1956 year class), 28.7% age II fish (1955 year class), 53.2% age III fish (1954 year class), and 7.6% age IV fish (1953 year class). Mayorova and Tkacheva (1959) noted that a rich year class in 1938 also dominated the fishery in the Black Sea from 1938 to 1945. The 1963 length-frequency distribution of S. sarda landed at Barbate and Tarifa, Spain, showed a strong mode centered at around 41 cm and lesser modes be- tween 49 and 59 cm and 59 and 69 cm (Fig. 21). The 1964 sample had only two modes: one at about 43 cm and the other at about 51 cm. Rodriguez-Roda (1966) also pre- sented a weight-frequency distribution of S, sarda (Fig. 22), which showed only two modes for the 1963 sample and only one well-defined mode for the 1964 sample. The 1964 data were also summarized by month (Fig. 23). The length-frequency distribution showed two well-defined modes in May, June, and July whereas the weight-fre- quency distribution showed only a single mode during those months. It is of interest that the relative propor- 20 iii 1963 - - - i i i i i i i — i \ \ i \ I \ . 1964 \ \ / \ 1 ^ - / ; / / / i / / / / / / J i / / \ V \ \ \ V \ \ \ \ \ \ \ \ ^ V v /"' — \ - ^■— T ,~^r^ "~ 35 40 45 50 55 60 65 70 LENGTH (cm) Figure 21. — Sarda sarda length-frequencv distribution. Barbate /,, + 2/„+/, 1Tl and Tarifa, Spain. Data smoothed by formula /„ = . (From Rodriguez-Roda 1966, fig. 11.) 4 tion of fish in the larger mode in the length-frequency distribution appeared to decrease from May to July. The 36 30 O 20- K A 1 1 1 1 1 /\ / \ - / \ \ A \ / \ ' / / / / / / ( / / -/ ' / / / \ / \ / \ / \ ; \ / \ 1963 \ \ \ \ .-1964 \ \ \ \ \ \ \ \ \ \ / / / / 7 / / \i i ■ \ V \ \ \ \ \ \ \ \ i - 12 3 4 5 WEIGHT (kg) Figure 22. — Sarda sarda weight-frequency distribution, Barbate /n-,+2/„+/„ + . and Tarifa, Spain. Data smoothed by formula f n = (From Rodriguez-Roda 1966, fig. 14.) 4 O 20- 40 50 LENGTH (cm) I 2 3 WEIGHT (kg) Figure 23. — Sarda sarda length (Barbate and Tarifa) and weight (Barbate) frequency distribution 1964. (From Rodriguez-Roda 1966, figs. 12, 15.) length-frequency distribution of male and female S. sarda were similar except that the modes were displaced (Fig. 24). 45 50 LENGTH (cm) Figure 24. — Sarda sarda length-frequency distribution, Barbate and Tarifa, Spain, arranged by sex. (From Rodriguez-Roda 1966, fig. 13.) Postel (1955a) presented data on the maximum size of S. sarda sampled in monthly periods in the eastern tropical Atlantic. The maximum lengths ranged from 450 to 690 mm for the males and 443 to 714 mm for the females. Rodriguez-Roda (1966) computed the length-weight relation of S. sarda landed at the southern Spanish port of Barbate (Table 23). 4.14 Subpopulations It appears that the S. c. lineolata found off northern Mexico and southern California comprise a single homo- geneous stock. Tagging experiments conducted by the California Department of Fish and Game indicate that the bonito does not make long migrations. Although some tagged fish have traveled as far as 600 miles, most of the tagged fish have been recaptured in the vicinity of release; fish that traveled long distances moved from Mexico to southern California waters in the summer and back to Mexican waters again in the winter (Frey 1971; Collins and MacCall 1977). From all indications it appears that the Northern and Southern Hemisphere populations of S. chiliensis are completely separate with little or no interchange. They are geographically separated from each other and there are certain meristic and morphological differences be- tween the two populations. For example, the Northern Hemisphere population of bonito averages more verte- brae than the Southern Hemisphere population (Vildoso 1963b; Kuo 1970; Collette and Chao 1975). Some preliminary work has been done on S. chiliensis to determine if protein differences attributable to genetic variation could be useful in identifying population units. Barrett and Williams (1967) experimented with 37 gel-electrophoresis of the soluble eye lens proteins of bonito in an attempt to find such genetically controlled differences. They did find polymorphisms of the soluble lens proteins for the bonito. They also calculated the gene frequencies together with their expected distribu- tions and found conformity to the Hardy-Weinberg prin- ciples. However, Barrett and Williams also found that the distributions of the apparent phenotypes were re- lated to the lengths of the bonito. They concluded, there- fore, that ontogenetic factors caused the observed poly- morphism and that gel-electrophoresis of the soluble eye lens proteins, under the conditions used in their experi- ment, was not a useful technique in differentiating popu- lation units of bonito. Smith (1971) examined the electrophoretic patterns of nuclear lens proteins from S. c. lineolata and also found polymorphism in the protein patterns. He suggested that there could be another explanation, as stated by Eckroat and Wright (1969), for the observed distribution of phenotypes than that suggested by Barrett and Williams (1967). Eckroat and Wright (1969) suggested that two separate fractions inherited as present/absence differ- ences could account for the distribution of the three phenotypes observed by Barrett and Williams (1967). If this situation obtained, then the distribution of the observed phenotypes would indicate that there were two different breeding populations represented in the catches sampled. It is not clear whether the stocks of S. c. chiliensis off Peru and Chile are independent of each other or whether they form a single homogeneous population. The center of bonito abundance off Peru is from Chimbote to Pisco (Ancieta 1963), which is centrally located between the north and south borders of Peru. The center of abun- dance off Chile is closer to the northern boundary of Chile near Arica, Iquique, and Antofagasta (Barrett 1971). In other words, the stocks of S. c. chiliensis off the 20 18 16 14 m 12 O X Q 8 8 6 <\ 2 — 1 — 1 1 1 X /\ / \ / \ / \ / / ANNUAL \ ^^ \ Y ^^"^ a. \ Y 3 o-~^ ,o -,:nov y° y ■•■•. y ^O^ \. s 2°- ■■-,>. ^-^--r~ S*' In "**■ ^ ^* v> -^- J ^-» ^C" .-° *•.. O"..^ ^o-- "y '"***"-o "■f 1965 1966 1967 1968 1969 Figure 25. — Apparent abundance of Sarda chiliensis chiliensis, for individual quarters and years, by year, from data for the bonito vessels. (From Barrett 1971, fig. X.) coasts of Chile and Peru are at least contiguous with each other. In his attempts at determining a stock production model for the Chilean bonito, however, Barrett (1971) apparently considered only the Chilean stock. He listed as one of his recommendations, however, that future studies should include a "determination of relation, if any, with the Peruvian fishery for bonito." No information is available on subpopulations of S. orientalis and S. sarda. The coastal distribution of all the species of Sarda, the migration patterns of S. c. lineo- lata, and the disjunct distribution of S. orientalis and S. sarda suggest the existence of discrete subpopulations. Based on scattered anatomical and meristic data, there appeared to be no difference in the populations of S. ori- entalis, although some morphometric differences were found in samples from Japan and the eastern Pacific. Similarly, a geographical comparison of S. sarda samples showed no conclusive differences among the various populations (Collette and Chao 1975). 4.2 Abundance and density Sarda australis Marshall (1964) said that S. australis, along the coasts of Queensland and New South Wales, are found in "great schools throughout the year." Grant (1972) stated that this species occurs in schools in Queensland waters espe- cially during the winter months. Sarda chiliensis In the Chilean bonito fishery Barrett (1971) examined the relations among fishing effort, yield, and apparent abundance for the period from January 1965 to Decem- ber 1969. Barrett recognized three different types of ves- sels, according to species objective, that fish bonito in the Chilean fishery: the anchovy, bonito, and tuna vessels. As the name implies, the anchovy and tuna vessels fished for bonito when anchovy and tunas, respectively, be- came less available and bonito more available. The bonito vessels fished primarily for bonito, although they did also capture other species. Barrett used the data from the bonito vessels for his analysis. The data for the entire monitored fleet, however, showed the same trends as for the bonito fleet after they were standardized to that of the bonito vessels. He noted a steep downward trend in total catch, an upward trend in relative fishing intensity, and a corresponding decline in apparent abundance in the fishery (Figs. 25, 26, 27). The trend appeared to level off in 1969. Barrett stated that the decline probably re- sulted from the effects of the fishery and not from fish- ery-independent factors. MacCall et al. (1976) related the indices of abundance of the Northern Hemisphere S. chiliensis obtained by Squire (1972) on aerial surveys with catch per unit of ef- fort (CPUE), in terms of catch in numbers of fish per angler, for the California party-boat fishery. MacCall et al. assumed that the aerial survey concentrated on the commercial fishing grounds because the surveys were 38 b(J 50 1 1 1 4 1 1 1 1 1 ! 1 I /\ t l\ TOTAL CATCH i 40 / \ \ l\ o \ \ 1 \ X 30 j \a / 1 / 1 o _l *: 20 ^J 1 M / 10 W \y i i — i — i — i — i — r RELATIVE FISHING INTENSITY 1 i 1 I 1 1 1 1 1 ! 1 1 1 1 1 1 1 1 1 A CATCH PER UNIT OF EFFORT t! i i b — S.J965 \ \. 1967 \ \ \ N \ N. V \ 1968 • o 1 1 l ; i i 18 19 20 21 22 23 24 25 26 27 28 RELATIVE FISHING INTENSITY ( DAYS X I0 2 ) Figure 27. — Apparent abundance of Sarda chiliensis chiliensis in relation to relative fishing intensity from data for the bonito vessels only (dashed line) and for the fleet monitored by IFOP (Instituto de Fomento Pesquero) (solid line), 1965-68. (From Barrett 1971, fig. 9.) 2 3 4 2 3 4 2 3 4 2 3 4 2 3 4 1965 1966 1967 1968 1969 X LU Q Z ol972 0.5 1.0 1.5 2.0 AERIAL SURVEY DAY INDEX Figure 28. — Regression of aerial survey day index against CPUE adjusted for mortality and recruitment for Sarda chiliensis lineo- lata. (Note: Although the authors did not state how the line was fit, presumably, the least squares method was used.) (From MacCall et al. 1976, fig. 12.) waters and became re-established as a locally spawning population." During the 1960's young S. c. lineolata were abundant inshore and sports fishermen landed record numbers; however, since 1969 the sport catch has fallen drastically (Collins and MacCall 1977). Sarda orientalis The mean monthly landings and catch rates for S. ori- entalis in waters off southern Japan (Fig. 30) showed that 39 g 1.5 < CPUE INDEX (WEIGHTED BY AREAS) IC/If 1935 1940 1945 1950 1955 I960 1965 1970 1975 YEAR Figure 29. — Party-boat Sarda chiliensis lineolata CPUE as an index of annual recruitment. (From MacCall et al. 1976, fig. 13.) 25 JAN FEB • MAR APR MAY JUNE JULY AUG SEPT. OCT NOV. DEC. Figure 30. — Seasonal landings and catch rates of Sarda orientalis in southern Japan. A — Mean monthly catch per trip by trolling in the Aburatsu, southern Kyushu, Japan, fishery for Sarda ori- entalis, 1970-73. B — Mean monthly landings of Sarda orientalis at Tosashimizu and Muroto, Kochi, Japan, by various fishing methods 1967-73. (Data provided by S. Kikawa, Far Seas Fish- eries research laboratory, and T. Koto, Nansei Regional Fisher- ies Research Laboratory, Japan.) ability or abundance of S. orientalis off the coast of southern Japan. Sarda sarda Wide fluctuations in the landings of S. sarda in the Black Sea-Mediterranean Sea area are a characteristic of this fishery (Fig. 31), and investigators in this area have been trying to find the causes of the fluctuations. Artuz (1959) noted that the landings at the Istanbul fish market showed fluctuations following a 9-yr cycle during the period from 1936 to 1958 (Fig. 32). As stated earlier, part of the fluctuations were explained by the entry into the fishery of rich year classes. However, Mayorova and Tkacheva (1959) pointed out that although relatively good year classes develop in certain years, these fish fail YEARS Figure 31. — The catches of Sarda sarda recorded at the Istanbul fish market, Turkey, 1909-60 (1 pair = two fish). (From Demir 1963, fig. 13.) > HYPOTHETICAL FLUCTUATION CURVE > SMOOTHED VALUES ACCORDING TO THE FORMULA i 3 2- - i i : ' ' ' ' ; 9""9 —VERY GOOD YEARS- l i / \ 1 \ 'R ' -f 1 1 1 1 1 1 I 1 1 1 1 I l\ 1 / \ 1 / V - / \ i / \ 1 ' i \\ f \ o— GOOD YEARS— f \ \ ' 1 1 1 \ \ S \ i 1 1 H \ - \ i 1 \ t ' V i i Vi A ' \\ /\ ' I \ \ 1 \ // // // \ \ L \ - V / \ ' 1 _MEDIUM V^>^/ \L-J YEARS \ 1 \ 1 1 / . / | / v \ \ \ \ ° \ I \ 1 \ 1 V\ \1 // // // // il if 1 \ 1 \ 1 \ 1 \ 1 —BAD YEARS — ' v J I 1 1 I 1 I I 1 I I r ^ti i i ! 1 1 1 both the landings and catch per trip of S. orientalis were relatively higher during the fall-winter months, indi- cating that there are seasonal differences in the avail- YEARS Figure 32. — Fluctuation pattern of Sarda sarda landed at the Istan- bul fish market 1936-58. (From Artuz 1959, fig. 1.) 40 to return to the Black Sea. They concluded that ". . . the rich yield of young fish in the Black Sea is not always fol- lowed by an increase in the abundance of large pelamid." Artuz (1959) also indicated a possibility of an inverse relation between the relative abundance of S. sarda and the mackerel, Scomber scombrus, in Turkish waters from 1940 to 1956 (Fig. 33). 1 1 1 1 1 1 1 1 1 1 1 1 Sarda sarda I0 6 PAIRS o ° K • \ I \ \ \ \ \ \ b \ s \ f \ / \ / \ / \ / \ / \ / \ / y ' \ / / i ; / / Uskumru 1 1 1 1 v°- **o— -o io 5 kg [III 1955 YEARS Figure 33. — Alternative occurrence of Sarda sarda (pairs) and uskumru, Scomber scombrus, (kg) landed from Turkish waters during the period 1940-56. (From Artuz 1959, fig. 5.) 4.5 Dynamics of population Barrett (1971) was unsuccessful in developing a stock production model, following Schaefer (1954), for the Chilean bonito fishery. He cited as the cause of this the fact that the necessary assumptions for this type of model were unsatisfied. One of the requisite as- sumptions of the model is a stable age and size distribu- tion in the population. The rapidly occurring changes in the fishery since 1965 have likely made this assumption untenable, according to Barrett. Another factor that af- fected the development of the model was the apparent discontinuous seasonal availability of some size classes of bonito in the fishery. Barrett concluded that he could not determine whether the fishery was stabilized or in a state of overfishing, and therefore recommended an acceler- ated research program on the Chilean bonico. MacCall et al. (1976) developed a surplus production model for the Northern Hemisphere bonito in California waters (Fig. 34). They used total catch data and a com- bination of CPUE for the party-boat fishery and aerial survey index for the period 1963 through 1972 to develop the model. The model indicated that the Northern Hem- isphere bonito is being harvested at or above the maxi- mum sustainable yield. MacCall et al. cautioned, however, that the assessment was confounded by the possibility of a density independent decline in recruit- ment. They noted that the California catch of S. c. lineo- lata in 1973 appeared to be greatly in excess of the equilibrium yield. They stated, however, that con- clusions drawn from the production model must be viewed with caution because bonito fishing in California waters is influenced by ocean temperatures. Because of the lack of necessary data, MacCall et al. did not attempt a yield per recruit analysis. Collins and MacCall (1977) determined a spawner- recruit relationship for S. c. lineolata during the period from 1964 to 1974 and found no simple functional curve to describe the relationship. They found that in some years a large spawning biomass was related to larger re- cruitment than did a small spawning biomass; however, in some years poor recruitment was also related to a high spawning biomass as well as to a low spawning bio- mass. 3 JO 15 20 25 30 35 EFFORT INDEX (XIO 7 ) 40 45 50 Figure 34. — Sarda chiliensis lineolata equilibrium yield and abun- dance in California waters. (From MacCall et al. 1976, fig. 14.) 4.6 The population in the community and the ecosystem MacCall et al. (1976) analyzed trophic interaction be- tween forage (northern anchovies, Engraulis mordax, and Pacific sardines, Sardinops sagax) and game fish (yellowtail, Seriola dorsalis; bonito, Sarda c. lineolata; albacore, Thunnus alalunga; barracuda, Sphyraena argentea; and white seabass, Cynoscion nobilis) in Cali- fornia waters. It appeared that there was little evidence that either forage or game fish biomass showed strong de- pendence on the abundance of the other. It appeared more likely that the abundance of both was determined largely by external conditions. 41 5 EXPLOITATION 5.1 Fishing equipment Sarda australis Sarda australis are generally taken incidentally by trollers who use the fillets as snapper bait or by sports fishers who use it as bait for billfishes and sharks (Grant 1972). Sarda chiliensis The California fishery uses various kinds of gear to catch S. c. lineolata including trolling gear and purse seine. The largest landings are made by the local purse seiners and the trolling fleet lands a lesser amount. The high-sea purse seiners may occasionally catch bonito to fill out the loads of other tunas (Frey 1971). In a study of the California sport fishery it was found that S. c. lineolata made up the biggest part of the fish catch by a single species (Pinkas et al. 1968). Bonito are taken in party and private boats, from piers and jetties, and from the shoreline. They will strike at most bait and lures, and fishing techniques for bonito vary from still fishing to trolling. The Chilean bonito fishery prior to 1964 was conducted almost entirely on an "artisanal, semi-industrial day- fishery basis" primarily from the ports of Iquique and Antofagasta (Barrett 1971). The vessels used in this fish- ery are known as "faluchos" and are about 10 m long. They fish principally with floating gill nets or with small purse seines called "boliches," which are hauled manual- ly. During the period from 1964 to 1966, two modern 36 m tuna/bonito seiners and eight other bonito/tuna seiners entered the fishery. All the new purse seiners in the bonito fishery have power launches and power blocks and use nylon nets ca. 549-732 m (300-400 fathoms) long and ca. 73-110 m (40-60 fathoms) deep. The fishing gear and vessels of the Peruvian fishery have been described by Ancieta (1963). Sarda orientalis In Japan this species is taken by various gear including trolling gear, pole and line, purse seines, and set nets. There is no exclusive fishery for S. orientalis in Japan; it is taken together with species that inhabit or enter the coastal waters of Japan in multispecies fisheries employ- ing various gear. In the Philippines S. orientalis are taken mostly in traps (Warfel 1950). In the Indian Ocean fishery S. orientalis are taken primarily by drift nets (gill nets) in India and Sri Lanka. Sarda sarda Sarda sarda are taken by trap net, ring net, gill net, trammel net, purse seine, beach seine, and hook and line (Demir 1963). Figure 35. — Fishing ports of the Peruvian and Chilean fish- eries for Sarda chiliensis chiliensis. (Adapted from Barrett 1971, fig. 1.) 5.2 Fishing areas Sarda australis There apparently is no well-developed fishery for S. australis in Australia. It occurs on the eastern coast of Australia along Queensland, New South Wales, and as far south as Port Fairy in Victoria. Serventy (1941a) stated that ". . . the bonito appears to be numerous enough to be regarded as a commercially important tuna, and a fishery could be maintained throughout the year on the eastern coast." Marshall (1964) noted that S. australis is destined to be of economic importance in Australia some day. There seems to be a difference of opinion on the food quality of 5. australis. Grant (1972) stated that the flesh of the Australian bonito is dark red and its edible qualities are not highly regarded, whereas Marshall (1964) claimed that the flesh is light-colored, of delicate flavor, and of good canning quality. Sarda chiliensis The Northern Hemisphere S. chiliensis is most abun- dant in the area from Point Conception, Calif., to Magdalena Bay, Baja California (Frey 1971), and pre- sumably this is the area in which the California-Mexico fishery for this species is conducted. Although the market for this species is limited, this bonito has been commercially fished in California waters since around 1900. However, the percentage of the California catch 42 made north of the California-Mexico border has been de- clining. From 1965 to 1968, over 90% of the California catch was made north of the border; during 1969-74, it was reduced to 70%. There are two semi-independent S. c. lineolata fisheries in the northeastern Pacific: a south- ern California fishery conducted by small purse seiners and a fishery conducted by the high-seas tuna fleet in southern Baja California (MacCall see footnote 5). The Southern Hemisphere population of S. chiliensis is exploited by the fisheries of Peru and Chile. The Peru- vian fishery extends along the coastline from the port of Mancora (ca. lat. 04°S, long. 81 °W) to the port of Ilo (ca. lat. 17°30'S, long. 71°15'W) (Ancieta 1963), and the Chilean fishery extends from Arica (ca. lat. 20°15'S, long. 70°15'W) to Talcahuano (ca. lat. 37°45'S, long. 73°W) (Barrett 1971) (Fig. 35). most abundant in the coastal waters of Kyushu (Kikawa and Staff of the Nankai Fisheries Research Laboratory, Kochi, Japan 1963). In the Indian Ocean minor fisheries for S. orientalis have been reported off the southwest tip of India (Silas 1964), in the Gulf of Aden along the coast of Somalia (Laevastu and Rosa 1963), and around Sri Lanka (Si- vasubramaniam 1969). 5.3 Fishing seasons Sarda australis No information. Sarda chiliensis Sarda orientalis Fisheries for this species are not very well developed throughout its distributional range. The FAO Yearbook of Fishery Statistics (e.g., FAO 1974) does not show any S. orientalis landings. The S. orientalis found in the tropical eastern Pacific between central Baja California and Peru is of little commercial importance and when caught may enter the catch statistics grouped with S. chiliensis (Pinkas 1961). This species is uncommon around Hawaii, and when landed the catches are not identified in the statistics published by the State. In Japan S. orientalis occurs in waters south of central Honshu along both the Pacific and Japan Sea coasts. It is i i — | i — . f— 1 | — 1 i — | 1 n Jl _ JAN. FEB MAR. APR MAY JUNE JULY AUG SEPT. OCT NOV. DEC. Figure 36.— Mean monthly landings and shipments into California of Sarda chiliensis lineolata, 1968-72. (Data from Heimann and Carlisle 1970; Pinkas 1970, 1974; Bell 1971; Oliphant and Marine Fisheries Statistics Staff. Marine Resources Region 1973.) The mean monthly landings of S. c. lineolata in Cali- fornia for the 5-yr period from 1968 to 1972 indicate that the fishery is seasonal (Fig. 36). Although bonito was landed throughout the year in this 5-yr period, peak landings occurred in August-October. Bonito is landed throughout the year in Chile and Peru, but both fisheries have seasonal peaks (Fig. 37). The seasonal peak is in October-November in Chile, whereas it occurs in January in the Peruvian fishery. 20 PERI 1 - - i— i - - — . _ CHILE 1 1 □ 1951,1958,19 O 1965-1968 59 8 1963 R i 'ft ; 1 (-1 r-fl 771 : . 1 1! :, 1 1 ft] JAN. FEB. MAR APR MAY JUNE JULY AUG SEPT OCT, NOV. OEC. Figure 37. — Mean monthly landings of Sarda chiliensis chiliensis in Peru, 1951-59 (data from Ancieta 1963, table 1) and Chile (from Barrett 1971, fig. 2.) 43 Sarda orientalis In the Indian fishery stray specimens may be taken in April. May, and June; however, the main fishing season appears to be from about July to September (Silas 1963). In nearby Sri Lanka, 5. orientalis appears in the catches throughout the year off the south coast. Mature fish are taken along the northeast coast from June-August and off the south and southwest coasts between September and February. The juveniles appear off the west coast usually during the southwest monsoon in June-August (Sivasubramaniam 1969). Sarda sarda The fishing seasons for S. sarda in various areas are summarized in Table 8. Rodriguez-Roda (1966) sum- marized the fishing seasons in the Spanish fishery by various geographical areas in the eastern Atlantic and the western Mediterranean (Table 9). Apparently the fishing seasons in the Black Sea- Aegean Sea areas are re- lated to the migrations of S. sarda. There is disagree- ment on the exact routes of the migration in certain areas, and the routes are unclear in other areas, but it is certain that S. sarda does migrate back and forth from the Aegean Sea, Sea of Marmara, and the Black Sea. Ac- cording to Demir (1957), schools of S. sarda spend the summer in the Black Sea and the winter in the Sea of Marmara and the Aegean Sea. Thus, the fishing season of May-October coincides with the time the migrating S. sarda are in the Black Sea in the summer. The fishing peaks of April-May and September-December in the Sea of Marmara coincide with the spring migration of the bonitos from the Aegean Sea through the Sea of Mar- mara to the Black Sea and the fall return migration from the Black Sea to the Aegean Sea. No information is available on the migration of S. sarda in the other areas of the Mediterranean and in the Atlantic Ocean. The seasonal development of the Spanish fishery in various areas of the western Medi- terranean, however, suggests that S. sarda may be mi- grating to and from the western Mediterranean (see Table 9). 5.4 Fishing operations and results The four species of Sarda are the bases of many fisher- ies throughout the world. Detailed information on the fisheries, however, is available only from a relatively few fisheries. Sarda australis, being found only along the coast of eastern Australia and around Norfolk Island east of Australia, is the object of a small incidental fishery in Australia. According to statistics published by FAO in the Yearbook of Fishery Statistics, S. c. lineolata is the basis of fisheries by Mexico and the United States. Sarda c. chiliensis is exploited by Chile and Peru. Landings of S. orientalis do not appear in the FAO Yearbook of Fishery Statistics (see FAO 1974). Relatively small amounts of the species are taken throughout its distri- butional range and it is not the basis of any well-de- Sarda sarda 1965 1966 1967 1968 1969 1970 1971 1972 1973 Figure 38. — Annual world landings of Sarda chiliensis and Sarda sarda, 1965-73. (Data from FAO 1974.) veloped fishery. Sarda sarda, on the other hand, appears to be exploited throughout its distributional range in fisheries of various sizes. In the Atlantic Ocean, the FAO Yearbook lists 18 countries reporting landings of S. sarda, mostly from the eastern Atlantic. Thirteen countries report landings of S. sarda from the Medi- terranean and Black Seas. The total landings of S. chili- ensis ranged from 53,300 to 94,100 t during the period 1965-73 and were greater than the total landings of S. sarda in each of 9 yr. The landings of S. sarda ranged from 25,000 to 65,700 t during the same period (Fig. 38) (FAO 1974). Sarda australis As can be expected from an incidental type fishery, the annual landings of S. australis were erratic and in some years no fish were landed at all during 1955-73 (Fig. 39). Information is meager on the utilization of the S. australis that are landed. They apparently are not un- common in the wholesale fish market of Sydney, Australia (Collette 6 ). Sarda chiliensis The landing figures of S. c. lineolata in California and Mexico show that Mexican landings make up only a small proportion of the total in the Northern Hemis- phere (Fig. 40). From 1965 to 1973, the California landings constituted from about 77 to 100 c "c of the total landings of bonito north of the equator in the eastern Pa- cific. The annual landings have been erratic, and to some ''Collette, B. B., Systematics Laboratory, National Marine Fisheries Service, NOAA, National Museum of Natural History, Washington, D.C. 20560, pers. commun. February 1978. 44 2 - — 1 . i i i i i r 1 Z2 ZJ " NO CATCH NO CATCH NO CATCH I [-] Zl ZZ\ NO CATCH NO CATCH NO CATCH III . . ID 113 IP (C tC m m m m in to en 0) en en 2.0 4.6 3.8 5.4 7.6 6.1 5.7 4.0 Total 1.9 2.4 5.1 4.6 6.3 8.4 7.1 6.9 5.8 Mediterranean and Black Seas: 26.9 22.1 41.1 25.3 55.2 10.6 22.9 21.9 22.0 The annual landings of S. sarda in the eastern and western Atlantic and in the Mediterranean -Black Seas during the period from 1965 to 1973 show that except in 1973 when the eastern Atlantic landings exceeded 10,000 t, the landings in the eastern and western Atlantic never rose above this figure (Fig. 43, Table 26). The fisheries of Spain and Portugal make up the landings in the eastern Atlantic. In the western Atlantic the bulk of the landings is made by the fisheries of Argentina and Brazil. The landings in the Mediterranean and Black Seas fluctu- ated from 10,600 to 55,200 t during the period from 1965 to 1973. Turkey contributed 71.7 to 91.3% of the annual landings during this period. 6 PROTECTION AND MANAGEMENT 6.1 Regulatory measures There are no measures regulating the commercial fish- eries for any of the species of Sarda. The State of Cali- fornia imposes a bag limit of 10 S. c. lineolata per day for recreational fishermen (Frey 1971), and is presently seek- ing to impose regulations on the California commercial fishery (MacCall see footnote 5). LITERATURE CITED ANCIETA, F. 1963. La pesqueria del bonito en el Peru. FAO Fish. Rep. 6, 3:1607-1619. 1964. Sinopsis sobre la biologia y pesqueria del "bonito" Sarda chilensis (Cuvier y Valenciennes) frente a la costa del Peru. Rev. Fac. Cienc. Biol., Univ. Nac. Trujillo l(l):17-49. ARTUZ, I. 1959. Fluctuations in the catches of some pelagic fishes in the Mar- mara and Black Seas. Poc. Tech. Pap. Gen. Fish. Counc. Medit., FAO 5:303-309. AUSTRALIA. FISHERIES BRANCH, DEPARTMENT OF PRI- MARY INDUSTRY. 1957. Fish. Newsl. 16(3), 31 p. 1958. Fish. Newsl. 17(3), 35 p. 1959. Fish. Newsl. 18(3), 35 p. 1960. Fish. Newsl. 19(3), 35 p. 1961. Fish. Newsl. 20(3), 35 p. 1962. Fish. Newsl. 21(3), 43 p. 1963. Fish. Newsl. 22(4), 43 p. 1964. Fish. Newsl. 23(3), 47 p. 1965. Aust. Fish. Newsl. 24(3), 46 p. 1966. Aust. Fish. Newsl. 25(3), 46 p. 1967. Aust. Fish. Newsl. 26(3), 48 p. 1968. Aust. Fish. Newsl. 27(3), 48 p. 1969. Aust. Fish. 28(3), 52 p. 1970. Aust. Fish. 29(3), 56 p. 1971. Aust. Fish. 30(3), 52 p. 1972. Aust. Fish. 31(3), 56 p. 1973. Aust. Fish. 32(3), 68 p. 1974. Aust. Fish. 33(5), 76 p. BARNHART, P. S. 1927. Pelagic fish eggs off La Jolla, California. Bull. Scripps Inst. Oceanogr., Univ. Calif. Tech. Ser. 1:91-92. BARRETT, I. 1971. Preliminary observations on the biology and fishery dynamics of the bonito {Sarda chiliensis) in Chilean waters. [In Engl, and Span.] Inst. Fom. Pesq. Bol. Cient. 15, 55 p. BARRETT, I., and A. A. WILLIAMS. 1965. Hemoglobin content of the blood of fifteen species of marine fishes. Calif. Fish Game 51:216-218. 1967. Soluble lens proteins of some scombroid fishes. Copeia 1967:468-471. BEAUCHAMP, J. J., and J. S. OLSON. 1973. Corrections for bias in regression estimates after logarithmic transformation. Ecology 54:1403-1407. BELL, R. R. 1971. California marine fish landings for 1970. Calif. Dep. Fish Game, Fish Bull. 154, 50 p. BELLOC, G. 1954. Les thons de la Meditterranee. Premiere note: germon, pela- mide et melva. Proc. Tech. Pap. Gen. Fish. Counc. Medit., FAO 2:283-318. BERG, L. S., A. S. BOGDANOV, N. I. KOZHIN, and T. S. RAFF. 1949. Tromyslovye ruby SSR (Commercial fishes of the US- SR). [In Russ.] Moscow, Pishchepromizdat, 787 p. BIGELOW, H. B., and W. C. SCHROEDER. 1953. Fishes of the Gulf of Maine. U.S. Fish Wildl. Serv., Fish. Bull. 53, 577 p. BOESEMAN, M. 1947. Revision of the fishes collected by Burger and von Siebold in Japan. Zool. Meded. (Leiden) 28:1-242. 1964. Scombroid types in the Leiden Museum collection. Proc. Symp. Scombroid Fishes. Mar. Biol. Assoc. India, Symp. Ser. 1:461-468. BOSCHUNG, H. T, JR. 1966. The occurrence of the common bonito, Sarda sarda, in the northern Gulf of Mexico. Trans. Am. Fish. Soc. 95:227-228. 47 BUEN. F. DE. 1930. Estados larvarios y juveniles de la Sarda sarda (Bloch). Inst. Esp. OeeanogT. Trabajos 3, 32 p. 1932. Formas ontogenicas de peces (Nota primera). Notas Resum. [In Span.] Inst. Esp. Oceanogr., Ser. 2, 57:1-38. 1958. Peces del suborden Scombroidei en aguas de Chile. Rev. Biol. Mar., Valparaiso 7:3-38. (Engl, transl. in files of Natl. Mar. Fish. Serv., NOAA, Honolulu, HI 96812.) CAMPBELL, G., and R A. COLLINS. 1975. The age and growth of the Pacific bonito, Sarda chiliensis, in the eastern North Pacific. Calif. Fish Game 61:181-200. COLLETTE. B. B., and L. N. CHAO. 1975. Systematics and morphology of the bonitos (Sarda) and their relatives (Scombridae, Sardini). Fish. Bull., U.S. 73:516-625. COLLINS, R. A., and A. D. MacCALL. 1977. California Pacific bonito resource, its status and manage- ment. Calif. Dep. Fish Game, Mar. Res. Tech. Rep. 35, 39 p. CUSHING, J. E. 1964. The blood groups of marine animals. Adv. Mar. Biol. 2:85- 131. DAWES, B. 1946. The Trematoda, with special reference to British and other European forms. Cambridge Univ. Press, 644 p. DE LA TOURRASSE, G. 1957. La peche au thon sur la cote basque poursuit sa modernisa- tion. Rev. Trav. Inst. Peches Marit. 21:561-592. DEMIR, M. 1957. Migrations of Sarda sarda Bloch in the Black, Marmara, and Aegean seas; the probable spawning places and times. Proc. Tech. Pap. Gen. Fish. Counc. Medit., FAO 4:127-134. 1963. Synopsis of biological data on bonito Sarda sarda (Bloch) 1793. FAO Fish. Rep. 6, 2:101-129. 1964. Distribution of meristic counts of common bonito (Sarda sarda Bl.) from Turkish waters. Proc. Tech. Pap. Gen. Fish. Counc. Medit., FAO 7:455-457. DEMIR, M., and N. DEMIR. 1961. Contribution to the knowledge of the eggs of the common bonito (Sarda sarda Bloch). Proc. Tech. Pap. Gen. Fish. Counc. Medit., FAO 6:213-218. DIEUZEIDE, R., M. NOVELLA, and J. ROLAND. 1955. Catalogue des poissons des cotes algeriennes. HI. Oste- opterygiens. Bull. Stn. Aquicult. Peche Castiglione, New Ser. 6, 384 p. DUCLERC, J., J. SACCHI, C. PICCINETTI, G. PICCINETTI- MANFRIN, A. DICENTA, and J.-M. BARROIS. 1973. Nouvelles donnees sur la reproduction du thon rouge (Thun- nus thynnus L.) et d'autres especes de Thonides en Medi- terranee. Rev. Trav. Inst. Peches Marit. 37:163-176. ECKROAT, L. R., and J. E. WRIGHT. 1969. Genetic analysis of soluble lens protein polymorphism in brook trout, Saluelinus fontinalis. Copeia 1969:466-473. EHRENBAUM, E. 1924. Scombriformes. Rep. Dan. Oceanogr. Exped. Medit. 1908- 1910, II (Biology) A, 11, 42 p. FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS. 1965. Catches and landings, 1965. FAO Yearb. Fish. Stat. 20, 360 P- 1970. Catches and landings, 1969. FAO Yearb. Fish. Stat. 28, var. pag. 1974. Catches and landings, 1973. FAO Yearb. Fish. Stat. 36, 590 P- FREY, H. W. (editor). 1971. California's living marine resources and their utiliza- tion. State Calif., Resour. Agency, Dep. Fish Game, 148 p. FURNESTIN, J., J. DARDIGNAC, C. MAURIN, A. VINCENT, R. COUPE, and H. BOUTIERE. 1958. Donnees nouvelles sur les poissons du Maroc atlanti- que. Rev. Trav. Inst. Peches Marit. 22:379-493. GNANAMUTTU, J. C. 1966. On the occurrence of the oriental bonito, Sarda orientalis (Temminck and Schlegel) along the Madras coast. J. Mar. Biol. Assoc. India 8:365. GODSIL, H. C. 1954. A descriptive study of certain tuna-like fishes. Calif. Dep. Fish Game, Fish Bull. 97, 185 p. 1955. A description of two species of bonito Sarda orientalis and S. chiliensis and a consideration of relationships within the genus. Calif. Dep. Fish Game, Fish Bull. 99, 43 p. GORBUNOVA, N. N. 1963. 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[In Russ. 50 FISHERIES SYNOPSES ; series of documents, issued by FAO, CSIRO, INP, and NMFS, contains comprehensive reviews of present knowledge on species and stocks of aquatic organisms of present or potential economic interest. The Fishery Resources and Environment Division of FAO is responsible for the overall coordination of the series. The primary purpose of this series is to make existing information readily available to fishery scientists according to a standard pattern, and by so doing also to draw attention to gaps in knowledge. It is hoped that synopses in this series will be useful to other scientists initiating investigations of the species con- cerned or of related ones, as a means of exchange of knowledge among those already working on the species, and as the basis for comparative study of fisheries resources. They will be brought up to date from time to time as further information becomes available. The documents of this series are issued under the following titles: FAO CSIRO INP NMFS Fisheries Synopsis No. Fisheries Synopsis No. Sinopsis sobre la Pesca No. Fisheries Synopsis No. Symbol FIR/S DFO/S INP/S NMFS/S Synopses in this series are compiled according to a standard outline described in Flb/S1 Rev. 1 (1965). FAO, CSIRO, INP, and NMFS are working to secure the cooperation of other organizations and of individual scientists in drafting synopses on species about which they have knowledge, and welcome offers of help in this task. Additions and corrections to synopses already issued will also be most welcome. 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