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Spawning Habitat of the Pacific Sardine (Sardinops Sagax) in the Gulf of California: Egg and Larval Distribution 1956-1 957 and 1971-1 991 M

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Spawning Habitat of the Pacific Sardine (Sardinops Sagax) in the Gulf of California: Egg and Larval Distribution 1956-1 957 and 1971-1 991 M HAMMANN ET AL.: PACIFIC SARDINE SPAWNING HABITAT IN THE GULF OF CALIFORNIA CalCOFl Rep., Vol. 39, 1998 SPAWNING HABITAT OF THE PACIFIC SARDINE (SARDINOPS SAGAX) IN THE GULF OF CALIFORNIA: EGG AND LARVAL DISTRIBUTION 1956-1 957 AND 1971-1 991 M. GREGORY HAMMANN' MANUEL 0.NEVAREZ-MART~NEZ Centro de Investigacibn Cientifica y Educacibn Superior de Eiisenada Instituto Nacional de Pesca Divisibn de Oceanologia Centro Regional de Investigaci6n Pecquera Km. 107 Cam. Tipana-Ensenada CaUe 20 No. 605 Sur Ensenada, Baja Cahfornia Guaymas, Sonora MCxico Mt-xico [email protected] YANIRA GREEN-RU~Z Graduate Ofice Instituto Nacional de Pesca Scripps Institution of Oceanography Crntro Regional de Invrstigacibn Pesquera University of California, San Diego Sibalo - Cemtos s/n. La Jolla, California 92093 Mazatlin, Sinaloa MCxico ABSTRACT separated localities on the Pacific Coast. Nevertheless, The distribution of Pacific sardine eggs and larvae in there may be regional dfferences in life-history traits and the Gulf of California was studied from 38 cruises car- population dynamics that occur on shorter time scales. ried out during 1956-57 and 1971-91. Eggs and larvae The Gulf of California is a semienclosed sea, unique were found throughout the gulf, but spawning was more in being the only large evaporation basin in the Pacific intense in the central region. Contrary to previous hy- Ocean (Roden and Groves 1959). It is characterized by potheses, spawning was not restricted to the coasts. Pacific great seasonality in temperature, circulation, winds, up- sardine spawn in the Gulf of California from November welling, and productivity (Rosas-Cota 1977; Badan- to May, most intensely during December and January. Dangon et al. 1985; Robles and Marinone 1987; Spawning habitat is inferred from the sea-surface tem- Va1dt.z-Holguin and Lara-Lara 1987; Bray 1988; Ripa perature (SST) where early-stage (1-111) eggs are present; and Marinone 1989; Alvarez-Borrego and Lara-Lara spawning occurred at 18.9" k 1.9"C. Water is rarely 1991; Paden et al. 1991; Cervantes-Duarte et al. 1993; colder than 14" in the gulf, so cold water is not likely Castro et al. 1994; Santamaria-del-Angel et al. 1994a, b; to limit spawning, but the probability of finding Pacific Lavin et al. 1995). sardine eggs is lower than 5% in waters warmer than 24" There have been many reports of sardine eggs and lar- at the surface. It is suggested that Pacific sardine spawn- vae collected from ichthyoplankton cruises in the Gulf ing in the Gulf of California is limited by the strong sea- of California (Sokolov and Wong-Rios 1972, 1973; sonality in sea-surface temperature caused in part by the De la Campa and Gutierrez 1974; Gutierrez 1974; summer intrusion of warm, subtropical water. Further- Gutierrez and Padilla 1974; Moser et al. 1974; Sokolov more, eggs and larvae appear to be retained by the cen- 1974; Wong-Rios 1974; De la Campa and Ortiz 1975; tral anticyclonic gyre found in the gulf during winter, Molina-Valdez and Pedrin 1975; De la Campa et al. and spawning close to the eastern coast could be detri- 1976; Padilla-Garcia 1976a, b, 1981; Olvera-Limas 1981; mental because of transport to the warm, subtropical Olguin et al. 1982; Molina-Valdez et al. 1984; Olvera- conditions in the south. Limas and Padilla 1986). However, egg stages have not been presented to better define the spawning season, and INTRODUCTION the overall seasonality of Pacific sardine spawning has not The Pacific sardine, Surdinops sugux (Jenyns, 1842), been described. is a coastal pelagic schooling fish found from the Gulf of The purpose of this paper is to analyze the seasonal- Cahfornia, Mexico, to British Columbia, Canada (Whte- ity in the distribution of Pacific sardine eggs and larvae head 1985). In the Gulf of Cahfornia, it has typically been collected on 38 cruises from 1956 to 1991 in the Gulf the dominant species in the multispecies "sardine" fish- of California and to determine the relation of spawn- eries. Yearly landings have exceeded 275,000 metric tons, ing to sea-surface temperature. Early-stage eggs are a and there is concern that the stock may have been over- good index of spawning, and their presence can be used exploited (Cisneros-Mata et al. 1995). Hedgecock et al. to define the thermal limits under which spawning tends (1989) found no genetic differences between sardine to occur (Tibby 1937; Ahlstrom, 1943; Lasker 1964; sampled &om the Gulf of Cahfornia and four other widely Lluch-Belda et al. 1991). MATERIALS AND METHODS 'Present address: Orbital Imaging Corporation. 21700 Atlantic Blvd., Dulles, Virginia 20166. [email protected] Historical raw data come from seven California [Manuscript received April 14, 1997.1 Cooperative Oceanic Fisheries Investigations (CalCOFI, 169 HAMMANN ET At.: PACIFIC SARDINE SPAWNING HABITAT IN THE GULF OF CALIFORNIA CalCOFl Rep., Vol. 39, 1998 I 33 TABLE 1 Cruises in the Gulf of California Number of Cruise Institution Dates stations CC5602 CXOFI, USA Feh. 1956 93 CC5604 Apr. 1956 129 CCj612 Ikc. 1956 79 CC5702 Feb. 1957 70 CC5704 Apr. 1957 12.5 CC5706 June 1957 132 CC5708 Aug. 1957 81 AA7101 INP, Mexico Apr. 1971 50 AH7110 Sept. 1971 80 AA7204 Apr. 1972 77 AH7206 Nov. 1972 79 AA7302 Mar. 1973 79 AH7303 Mar. 1973 15 AA7305 Apr.-May 1973 75 AA7308 Jdy 1973 103 AA7402 Feb.-Mar. 1974 52 AA7403 Apr. 1974 12 AA7405 Dec. 1974 19 AA7501 Jan. 1975 66 AA7503 Mar.-Apr. 1975 50 -- 23 AA7504 Apr. 1975 39 AA7601 Jan. 1976 6.5 7-22 AH7605 Apr. 1976 66 AA760.5 58 117 116 115 114 113 112 111 110 109 108 107 July 1976 AA7701 Feb. 1977 24 Figure 1. The Gulf of California. AH7703 Srpt. 1977 58 AA7704 Aug.-Srpt. 1977 50 AA7708 Dec. 1977 48 AA7802 Feb.-Mar. 1978 59 USA) and 28 Instituto Nacional de Pesca (INP, Mexico) AA7810 Sept.-Oct. 1978 59 AA8103 Mar.-Apr. 1981 78 cruises in the Gulf of California (fig. 1) during 1956-57 PU8403 Mar.-Apr. 1984 87 and 1971-87; three additional cruises were carried out PU8611 Nov. 1986 187 by the Centro de Investigacibn Cientifica y de Educacibn AA8701 Jan.-Feb. 1987 56 PU87 1 1 Nov. 1987 22 Superior de Ensenada (CICESE, Mexico) during 1990 AL9002 CICESE Feh.-Mar. 2990 200 and 1991 (table 1). BIPXI9008 Aug.-Srpt. 1990 38 The number of cruises per year, number of stations, PU9109 Sept. 1991 110 sampling gear, and station plan varied among cruises (fig. 2, table 2); there were no cruises during the 1960s. During the 38 cruises, 3,631 tows from 2,667 stations number of myomeres and pigmentation of the larvae were made with a surface neuston net, a standard CalCOFI have been reported (Watson and Sandknop 1996). net, a bongo net, or a CalVET net, following ichthy- Nevertheless, it is unlikely that thread herring eggs and oplankton techniques described by Kramer et al. (1972), larvae occurring in the summer would have been Smith and Richardson (1977). and Lasker (1985). misidentified as Pacific sardine for several reasons. Adult Pacific sardine eggs and larvae were separated; eggs distribution of Pacific sardine during the summer is re- were staged following Ahlstrom (1943); and larvae were stricted to the region of the large islands, north of the measured to k0.5 mm. Egg abundance data were stan- central gulf (Cisneros-Mata et al. 1988). Adult matu- dardized to numbers/m2; larval abundances were stan- rity studies have shown that Pacific sardine preferentially dardlzed to nunibers/10 m2. The high variabhty ofwater spawn in cooler temperatures during winter and spring volume filtered in the surface neuston tows caused by (Torres-Villegas et al. 1986), and thread herring spawn the net breaking the surface, and the overestimation of in warm temperatures during summer (Torres-Villegas abundance from surface samples does not allow for stan- et al. 1985; Rodriguez 1987). The relative abundance dardization, so for neuston samples we used presence of thread herring adults in the Gulf of California is usu- or absence, not actual abundance. ally lower than that of Pacific sardine (Cisneros-Mata et During summer months in the Gulf of California, al. 1988). As an example, thread herring scales were not Pacific sardine eggs could be confused with those of sufficiently abundant to appear in laminated sediments thread herring (Opisthonema spp.; Matus-Nivbn et al. in the Gulf of California (Holmgren and Baumgartner 1989 [for 0. libertate]),but slight differences between the 1993), even though they are quite different from Pacific 1 70 HAMMANN ET AL.: PACIFIC SARDINE SPAWNING HABITAT IN THE GULF OF CALIFORNIA CalCOFl Rep., Vol. 39, 1998 1992 TABLE 2 Number of Stations and Tows per Cruise in the 1990 Gulf of California 1988 Cruise Stations Oblique tows* Neuston tows CC5602 93 93 0 1986 CC5604 129 129 0 1984 CC5612 79 79 0 CCj702 7 0 70 0 1982. CC.5704 125 125 0 CC5706 132 132 0 AA7101 50 50B 50 AH71 10 80 80C 80 1978 AA7204 77 77c 77 AH7206 79 79c 79 1976 AH7302 79 71C 61 AA7303 15 15C 4 1974 AA7305 75 73c 70 AA7308 103 66C 103 1972 AA7402 52 40C 52 "47403 12 8C 12 1970 AA7405 19 16C 17 1968 AA7501 66 37c 66 AA7503 50 50C 50 1966 AA7504 39 32C 39 IIIl1IIIII AA7601 65 39C 65 1964 AH7605 66 49c 66 AA760.5 58 39c 58 1962 AA7701 24 21c 24 AH7703 58 48C 58 1960 AA7704 50 iOC 0 AA7708 48 48C 0 1958 AA7802 59 59C 0 AA7810 59 59c 0 1956 AA8103 78 78C 0 PU8403 87 87B 0 1954 IIIIIIIIIII IIIIIIIIIIII PU8611 187 187CV 0 1 2 3 4 5 6 7 8 9 101112 AA8701 56 %I) 0 Month PU8711 22 22u 0 AL9002 200 200cv 0 Fiaure 2.
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