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The Early Life History of Fish Rapp. P.-v. Réun. Cons. int. Explor. Mer, 191: 63-69. 1989 Coastal upwelling and the early life history of sardines {Sardina pilchardus) along the Galician coast of Spain Edward J. Chesney and Manuel Alonso-Noval Chesney, Edward J., and Alonso-Noval. Manuel. 1989. Coastal upwelling and the early life history of sardines (Sardina pilchardus) along the Galician coast of Spain. - Rapp. P.-v. Réun. Cons. Int. Explor. Mer, 191: 63-69. The oceanography of the Galician coast of NW Spain from Ria de Vigo north to Cape Ortegal was studied on four cruises. Conditions favoring upwelling dominated three of the four cruises while conditions favoring downwelling prevailed during the other cruise. Distributions of fish eggs and larvae along with length frequencies of sardine larvae Sardina pilchardus, suggested that, during conditions favoring upwelling, larvae tended to drift south and offshore south of Cape Finisterre, while off Cape Ortegal in the north larvae tended to move offshore to the west and north. During conditions favoring downwelling sardine eggs remained close to shore where they were spawned. Sardine eggs were generally associated with areas of strongest coastal upwelling, indicating a preference for these areas by adult sardines. Size frequency distributions of sardine larvae showed a significantly greater frequency of larger older larvae during the upwelling cruises than during the spring peak time (March/April) for sardine spawning. Densities of fish larvae were generally highest along the transect NW of Cape Finisterre at the corner of the Iberian peninsula when compared to densities observed on the Coruna and Arosa transects. The pattern suggested that the frontal convergence known to exist in that area was aggregating organisms offshore. The observed distributional patterns of sardine eggs and larvae were consistent with both the known and theoretical patterns of oceanographic flow as well as with the known distribution of adult sardines in the area. The potential significance of the oceanography and the relationship between upwelling and regional recruitment of Sardina pilchardus is discussed. Edward J. Chesney: Louisiana Universities Marine Consortium, Star Route Box 541, Chauvin, Louisiana 70344, USA. Manuel Alonso-Noval: Instituto Espanol de Oceanografia, La Coruna, Spain. Introduction distinguishes it from the rest of this coastal upwelling The NW corner of the Iberian peninsula is an interesting system is the convergence of two subsurface water study area from both an oceanographic and fisheries masses, North Atlantic Central Water (NACW) from perspective (Fig. 1). The area from Cape Ortegal south the west and modified NACW from the north (Fraga, to the Portuguese border is part of a general coastal 1981; Fraga et al., 1982; Estrada, 1984). upwelling system extending southward to approximately One of the prominent fisheries of the area is a coastal 15°N (Wooster et al., 1976; Fraga, 1981 ; McClain et al., purse seine fishery for sardines, Sardina pilchardus, 1986). The upwelling season is from March through which typically captures in excess of 100,000 tonnes October, but is most intense from April through August. annually along the Galician and Cantabrian coasts. The Galician coastal region is a unique part of this major Other regionally important pelagic fisheries are the upwelling system because the corner of the Iberian scad, Trachurus trachurus, and the anchovy, Engraulis peninsula allows conditions favoring upwelling to pre­ encrasicholus. The Galician coast is important to the vail through 270° of wind direction (McClain et al., regional sardine fishery because a large portion of the 1986). The unique coastal embayments south of Cape Iberian stock of sardines has been known to reside Finisterre also add considerable complexity and diver­ within the Galician upwelling zone during the summer sity to both the oceanography and biological pro­ upwelling season (Dias etal., 1983; Pastor et al., 1985a, ductivity of the region (Tenore et al., 1982; Blanton et b; Pastor et al., 1986a, b). Sardina pilchardus is well al., 1984). A third unique feature of this area which known for its association with and apparent preference 63 r so plankton (egg production, biomass, macro- and micro- P i Larvae zooplankton species composition), ichthyoplankton ■ Eggs (species composition, density), and bacteria (produc­ Scale = per 100 m3 0 tion, density). A comprehensive description of these . Cabo Ortegal studies will be presented elsewhere. Only a brief syn­ opsis of the physical conditions encountered during the cruises and results from the ichthyoplankton studies, LA CORUNA emphasizing Sardina pilchardus, will be presented here. Ichthyoplankton and macrozooplankton were 'Cabo Villano sampled with a paired opening-closing net frame (60 cm ^Cabo Finisterre diameter) fitted with 333 and 202 mesh nets. All Spain sampling was carried out during daylight hours. Volume filtered during each tow was estimated using a General Ria de Muros Oceanics flowmeter. Oblique tows were taken by low­ 00 ’RIAS BAJAS' ering the net closed to a target depth, opening the net -0 by messenger, and retrieving it at a constant speed of approximately 3 knots over a 10 min interval. The net m Ria de Pontevedra was closed with a second messenger at the upper target depth. Two depth intervals were sampled at each sta­ r**R ia de Vigo sO tion. On the first two cruises (June 1984 and September 25 50 1984) samples were taken from the surface to 50 and VIGO km 50-100 m, except at the innermost station where the Figure 1. Densities of sardine, Sardina pilchardus, eggs and fishable water column was divided evenly. On sub­ larvae along the Galician coast, NW Spain, June 1984. Values sequent cruises sample depths were determined by the are means of two depth strata (0-50 m, 50-100 m). depth of the chlorophyll maximum or mixed layer, generally 0-30, 30-60 m or 0-40, 40-80 m. Each sample was preserved in 5% formalin for later analysis. Fish eggs and larvae were sorted, identified to species when for coastal upwelling areas (Blackburn and Nellen, possible, and counted. All Sardina pilchardus larvae 1976; Pastor et al., 1985a, b, 1986a, b; Cabanas et al., were measured to the nearest 0.1 mm standard length 1985). for the length-frequency analysis. Several studies have reported on the early life stages The sampling strategy on the first two cruises was of fishes along the Atlantic portion of the Iberian pen­ different from that on cruises 3 and 4. On the first two insula (Dicenta etal., 1977; Suau and Vives, 1979; Re, cruises each station was sampled only once to get a 1981; Franco and Sola, 1985; Sola and Franco, 1985). spatial distribution (Fig. 1). For the third cruise transects Two of them have focused on fish eggs and larvae within off Ria de Arosa and North of Cape Finisterre were the rias (estuaries) of the Galician coast (Lopez-Jamar, sampled many times to study temporal responses to 1977; Ferreiro and Labarta, 1988a, b), but only one upwelling. The strategy was similar for the fourth cruise study has reported on the distribution of fish eggs and except that a transect off La Coruna was added. larvae on the Galician shelf (Sola and Franco, 1985). A composite figure of spawning activity along the This paper will focus on the distribution and abun­ Iberian peninsula was developed by taking egg abun­ dance of sardine eggs and larvae encountered during dance data from four sources: Re (1981), Dicenta the four cruises to study the oceanography of the region, (1984), Ferreiro and Labarta (1984), and Sola (1987). emphasizing the distributions of Sardina pilchardus Their data were normalized to a value of 100 for the early life stages in relation to the oceanography. highest monthly abundance observed during their annual studies and the monthly values averaged for all four studies to produce an index indicative of sardine Materials and methods spawning activity for the entire Iberian peninsula. The oceanography and associated biological pro­ ductivity of the Galician shelf were studied aboard the R/V “Cornide de Saavedra” on 16-26 June 1984, 27 Results September to 2 October 1984, 27 July to 8 August Synopsis of physical oceanography 1985, and 22 March to 2 April 1986. Several physical, chemical, and biological parameters measured during Cruise (16-26 June 1984). Upwelling-favorable winds the cruises included water column hydrography (tem­ (up to 13ms 1 from the NE) prevailed from 7 June, perature, salinity, nutrients, oxygen), phytoplankton increasing steadily in strength until 22-23 June when (primary production, chlorophyll, irradiance), zoo­ they subsided. The average daily upwelling index for 64 the dates of the cruise was +1390 m3 s“ 1 km-2. Satellite dae (15.4%), Callionymidae (8.8%), the flounder, surface temperature showed upwelling localized off Arnoglossus laterna (4.9%), Labridae (3.8%), the horse Cape Ortegal and Cape Finisterre with little upwelling mackerel, Trachurus trachurus (3.8%), and anchovy, visible at the surface in the south off the Rias Bajas. Engraulis encrasicholus (1.3%). Mean abundances of Subsurface temperature and nutrient data suggested a fish larvae during the four cruises were 29.2 100 m “3 in slightly different interpretation. Uplifted isotherms and June 1984, 19.8 100 m “3 in July 1985, 34.7 100 m '3 subsurface nitrate maxima were seen off Rias Arosa in March 1986, and 3.5 100 m '3 in September 1984 and Vigo indicating subsurface upwelling in the south. reflecting the high and low periods of annual spawning Density distributions indicated southward currents off activity in the area. the Rias Baj as. The sardine is the most abundant fish along the Gali­ cian shelf and because it is a serial spawner its eggs and Cruise (27 September to 2 October 1984). Winds were larvae are abundant and occur during most of the year. weakly downwelling-favorable, varying during the Distributions and abundance of sardine eggs and larvae beginning of the cruise from SSW shifting to WNW varied in response to season and oceanographic con­ toward the end of the cruise to a weak upwelling- ditions.
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